Bull. nat. Hist. Mus. Lond. (Zool.) 63(2): 93-122

The lucinid bivalve genus Cardiolucina | wis

(Mollusca, Bivalvia, Lucinidae): systematics,

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Issued 28 November 1997

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JOHN D. TAYLOR. AND EMILY A. GLOVER Pe Ny r

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK abe

CONTENTS

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SYNOPSIS. The marine lucinid genus Cardiolucina (usually known as Bellucina) comprises 11 living species of small, heavily

ornamented, subspheroidal bivalves. The type species of the genus is the Miocene fossil C. agassizi (Michelotti). A new

definition and description of the genus is given. All the living species and the type species are redescribed and illustrated. Two

species, C. australopilula from Western Australia and C. siquijorensis from the Philippines, are described as new. Anatomical

details are given for two species which both contain chemosymbiotic bacteria in the ctenidia. A preliminary phylogenetic

analysis using shell characters and Parvilucina as an outgroup, indicates three clades of Cardiolucina. Nine species are found

in the Indo-W. Pacific and two in the tropical Atlantic. The greatest diversity occurs around Australia where seven species live.

The two Atlantic species are similar to the widespread Indo-Pacific species C. semperiana. The earliest-known species of

Cardiolucina is from the Eocene, but the fossil record is very poor.

INTRODUCTION

Since the discovery of endosymbiotic sulphur-oxidising bacteria

contained in the gills, the biology of bivalves in the family

Lucinidae has attracted much attention (Fisher & Hand, 1984;

Dando, Southward, Southward, Terwilliger & Terwilliger, 1985;

Dando, Southward & Southward, 1986: Southward, 1986; Reid &

Brand, 1986; Distel and Felbeck, 1987; Cary, Vetter & Felbeck,

1989; Fisher, 1990; Reid, 1990). The symbiosis has so far been

recorded in about 25 species of Lucinidae and some species have

been shown to be nutritionally dependent on the association (Le

Pennec, Beninger & Herry, 1995). The discovery and investigation

of the symbiosis has been the key to understanding the many

unusual features of lucinoid morphology and habitat distribution

(Allen, 1958; Reid & Brand, 1986, Reid, 1990). Despite the bur-

geoning biological interest in lucinid bivalves, it is unfortunate

that the systematics of family are not well understood, with many

taxonomic groups, particularly at the supraspecific level, being

rather loosely defined. Most recent comprehensive systematic

treatments of Lucinidae (Chavan, 1969; Bretsky, 1976) both pre-

date the discovery of chemoautotrophy in the family and,

moreover, are based entirely on shell characters As Hickman

(1994) points out, there has been no attempt to incorporate the

potential wealth of anatomical and biological characters into sys-

tematic studies. Furthermore, following closer attention to lucinid

biology and the investigation of new habitats it is now being

recognised that there are many undescribed species.

In this paper, we review the species of the lucinid genus

Cardiolucina (better known as Bellucina), which are largely tropi-

cal, small, subspheroidal bivalves found in the Indo-Pacific and

Atlantic Oceans. The impetus for this revision stems from the

discovery of two coexisting lucinids in the shallow waters around the

Abrolhos Islands, Western Australia (Glover & Taylor, 1997). Both

species appeared to be undescribed and one with unusual periostracal

extensions has been assigned to a new genus Rastafaria (Taylor &

Glover, 1997). The other species, which forms a part of this paper,

was evidently a Cardiolucina and seemed similar to C. semperiana

Issel, which is widespread in the Indo-West Pacific. However,

preliminary research indicated that there is much taxonomic confu-

sion within the genus, including the priority of the name Cardiolucina

over Bellucina. This led us to revise and redescribe the type species,

provide a new diagnosis of the genus, to review the Recent species

from the Indo-Pacific and Atlantic Oceans and to describe new

species from Western Australia and the Philippines. We have also

attempted to identify within the genus Cardiolucina those shell and

anatomical characters that might be used in a wider study of lucinid

systematics.

ABBREVIATIONS

Institutions: AM — The Australian Museum, Sydney; ANSP —

Academy of Natural Sciences, Philadelphia; BMNH — The Natural

History Museum, London; MNHN — Muséum National d’ Histoire

Naturelle, Paris; IRSNB — Royal Belgian Institute of Natural Sci-

ences; NM — Natal Museum; NMW — National Museum of Wales,

Cardiff; NSMT — National Science Museum, Tokyo; SAM — South

Australian Museum, Adelaide; USNM — National Museum of Natu-

ral History, Washington; UMT — University Museum, Tokyo; WAM

— Western Australian Museum, Perth; ZMA — Zoological Museum,

Amsterdam; ZMC — Zoological Museum, Copenhagen.

Shell measurements: H= height of shell from umbone to ventral

margin, L = length of shell from anterior to posterior, T= tumidity of

shell i.e. maximum convexity measured on a single valve.

SYSTEMATIC DESCRIPTIONS

Family LUCINIDAE Fleming, 1828

Genus CARDIOLUCINA Sacco, 1901

Cardiolucina Sacco, 1901: 89. Published June 30 (see Sacco, 1904;

Marshall, 1991).

Bellucina Dall, 1901: 806. Published August 22nd (see Keen, 1971:

983). Lucina eucosmia Dall, 1901: 806, original designation.

TYPE SPECIES.

nation).

Cardium agassizi Michelotti, 1839 (original desig-

NOMENCLATURE. The species considered in this paper have usu-

ally been assigned to the genus Bellucina Dall, 1901. However, the

Miocene C. agassizi Michelotti, the type species of the genus

Cardiolucina Sacco 1901, has all the characters of Recent Bellu-

cina species (see description and figures of C. agassizi below).

Chavan (1937) recognised this and synonymized the two genera

giving Bellucina priority. Although published in the same year,

Sacco’s work appeared in June 1901 (Sacco, 1904, Marshall, 1991)

and Dall’s in August (see Keen, 1971). Cardiolucina therefore has

priority over Bellucina.

Because the name Bellucina has been commonly used, the con-

cept of this genus has to be considered in some detail. Dall (1901, p.

806) erected Bellucina as a section of the subgenus Parvilucina

within the genus Phacoides. The genus Parvilucina was originally

diagnosed as ‘Shell small, plump, often inequilateral; sculpture

more or less reticulate but not muricate, teeth small, common but all

usually present’ and the Section Bellucina was rather skimpily

distinguished as ‘Dorsal areas and sculpture strong.’ However, Dall

(1901) clearly designated Lucina eucosmia (= L. pisum Reeve) as

the type species and the concept of Bellucina must be based upon

this species. Many authors (e.g. Fischer, 1871, Lamy, 1920; Chavan,

1969; Britton 1972; Bretsky, 1976; Oliver, 1992) have considered B.

eucosmia to be synonymous with Bellucina semperiana (Issel,

1869). Consequently, the characters of the latter species have been

taken to represent Bellucina. Unfortunately, Reeve (1850) mixed

two distinct species in the original description and illustrations of

Lucina pisum which is clear from the syntype material. Reeve’s

figure 66a illustrates a specimen from Port Essington, Northern

Territory, Australia, and figure 66b shows a shell of another species

from Singapore. In the original description of Lucina pisum Reeve

clearly referred to the deeply incised lunule and fenestrate ornament

J.D. TAYLOR AND E.A. GLOVER

which are features of the Port Essington specimens. ‘Shell globosely

cordate, solid, thickly latticed with concentric and radiating ribs, of

which the interstices are pitted, posterior side grooved, anterior

short, lunule small, deeply excavated; whitish.’ For this reason one of

the syntypes from Port Essington has been selected as the lectotype

of Lucina pisum Reeve (see below under description of Cardiolucina

eucosmia). Because the name L. pisum is preoccupied (L. pisum

Sowerby, 1836) then C. eucosmia (Dall, 1901) becomes the valid

replacement name. The other Reeve syntypes from Singapore we

consider to be Cardiolucina semperiana (see below).

DIAGNOSIS. Shells small but solid, usually less than 14mm in

height, subcircular (Height/Length ratio around 1.0), moderately

inflated to subspheroidal (Tumidity/Length ratio between 0.38 and

0.48). Umbones prosogyrate, more or less central. Anterior sulcus

either absent or shallow and narrow. Posterior sulcus always

present and either deeply-indented or shallow. Lunule varies in

size and depth between species, but where visible it is generally

heart shaped. It can be deeply incised and large as in C. eucosmia,

or very small and hidden under the beaks as in C. australopilula.

Posterior dorsal area distinct, either concave or convex, usually

with concentric ribbing. Ornament of radial ribs and concentric

lamellae, relative strength of either varies between species. Con-

centric lamellae often raised and fluted. Intersection with radial

ribs produces a variety of ornament from fenestrate to beaded.

Hinge plate thick, dentition of single anterior and posterior lateral

and two cardinal teethin each valve. In the right valve of some

species one of the cardinal teeth can be either reduced or absent.

Ligament external and varying in length; can extend from beaks to

position of posterior lateral tooth, or may be two thirds of this

distance. Internal ventral margin finely to coarsely crenulate, cor-

responding to position of radial ribs. Often small denticles around

antero- and postero-dorsal margins. Anterior adductor muscle scar

quadrate to elongate, ventral tip only slightly detached from pallial

line. Mature individuals of spheroidal species often have the ad-

ductor muscle scars located on an internal buttress. Posterior

adductor scar short and rounded.

Juvenile shells are thinner, less inflated, more anteriorly extended

with radial ribs often visible from interior of shell. Interior ventral

margin not thickened.

ANATOMY. Based on observations of C. australopilula and C.

semperiana.

Mantle largely unfused except at posterior exhalant aperture.

Mantle margin thick and muscular. Mantle ‘gills’ absent. No fused

inhalant aperture. Exhalant aperture a muscular tube with flared

distal end, which can presumably be protracted. Ctenidia consist of

inner demibranchs only, filaments elongate and thick. Outer lamella

of demibranchs attached to posterior mantle just ventral to the

exhalent aperture. Labial palps are small ridges at edge of lips. Foot

cylindrical and vermiform with a posterior heel. Body wall muscu-

lar, with large hemispherical visceral pouch containing oocytes.

Rectum curves around dorsal side of the posterior adductor muscle

and opens near the posterior aperture.

Many of the anatomical features, such as the thickened inner

demibranchs, reduced labial palps and vermiform foot are similar to

those of other lucinids, although some characters may be less widely

distributed. A distinctive and possibly functionally important feature

is the attachment of the ctenidia to the posterior mantle. It is

uncertain how widely this character is distributed amongst the

Lucinidae. It was first illustrated by Reid & Brand (1986) for

Parvilucina tenuisculpta, but not mentioned by Allen (1958) in his

general survey of lucinoids, or by Morton (1979) for Fimbria and not

present in Rastafaria (Taylor & Glover, 1997). Large hemispherical

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

visceral lobes or pouches are not present in most Lucinidae (Allen,

1958), but are found in Parvilucina (Reid & Brand, 1986), and

species of Thyasiridae have several pouches (Allen, 1958). Most

Lucinidae seem to possess both inhalant and exhalant posterior

apertures, produced by mantle fusion (Allen, 1958). In Cardiolucina

there is no inhalant aperture, a state found also in Parvilucina (Reid

& Brand, 1986).

Cardiolucina species share three anatomical characters with

Parvilucina tenuisculpta, namely; the attachment of the posterior

ctenidia to the mantle, the hemispherical visceral pouches and the

lack of inhalant apertures. These characters may indicate a possible

relationship, but the distribution of most anatomical characters

amongst the Lucinidae is not well known.

GEOLOGICAL RANGE. Eocene (Lutetian) to Recent.

COMPARISON WITH OTHER GENERA. There has been a tendency to

place small, inflated lucinids with both radial and concentric orna-

ment, rather arbitrarily into a number of genera such as Phacoides,

Parvilucina, Linga and Bellucina. These decisions have usually

been made without reference to the characters of the type species.

For instance, the status of Bellucina in major taxonomic revisions

has varied as follows:

Phacoides (Parvilucina) section Bellucina — Dall, 1901: 806

Phacoides (Bellucina) — Lamy, 1920: 211

Linga (Bellucina) — Chavan, 1937: 205; 1969: 496

Parvilucina (Bellucina) — Britton, 1972: 15

Lucina (Bellucina) — Bretsky, 1976: 272-3

Cardiolucina (as Bellucina) has often been considered as a sub-

genus of Parvilucina Dall, 1901, type species Lucina tenuisculpta

Carpenter, 1864, which ranges from Alaska to Baja California.

Hickman (1994) redescribed and illustrated P. tenuisculpta and two

other west American species. Although they are small, rounded and

fairly inflated, Parvilucina species are thinner shelled and less tumid

(T/L 0.29) than all Cardiolucina (T/L 0.38-48) species. Most

Cardiolucina have a pronounced posterior sulcus, and sometimes a

narrow anterior sulcus, compared to the weak, shallow, posterior

sulcus of Parvilucina. Parvilucina has both radial and concentric

ribs; the radial ribbing consisting of closely-spaced, low riblets,

which are generally weak, but most prominent at the anterior and

posterior of the shell. The internal ventral margin of Parvilucina is

finely denticulate, compared with the coarser crenulations of

Cardiolucina. The lunule in Parvilucina is narrowly lanceolate and

asymmetric with the right valve overlapping the left, compared with

the shorter, heart-shaped lunule of Cardiolucina agassizi. The hinge

plate of Parvilucina is also thinner and the lateral teeth distant from

the cardinals. Although a major reappraisal of the relationships of

these genera is necessary, evidence suggests that the Parvilucina

taxa centred around the type species, represent a clade distinct from

Cardiolucina.

Another genus which should be considered in relation to

Cardiolucina is Radiolucina Britton, 1972, (type species Phacoides

amiantus Dall, 1901) from the Caribbean. Dall placed this species

and the similar West American species L. cancellaris Philippi, 1846

in Bellucina. Later, Keen (1971, p.121) recognised that they were

only superficially similar to the type species of Bellucina and

placed them in the subgenus Pleurolucina Dall, 1901, remarking

that American authors had been slow to recognise Dall’s error.

Bretsky (1976) confusingly gave an extensive diagnosis of the

genus Bellucina based on P. amiantus rather than the type species

Bellucina eucosmia. Britton (1972) had previously compared P.

amiantus with Bellucina semperiana (Issel) which he considered

the type species of Bellucina and recognised that P amiantus

represents a distinct and separate lineage within the Lucinidae. For

this reason, he erected a new subgenus Radiolucina. The type

species, R. amianta has a more elongate shell than most Bellucina

species H/L 0.92 (data from Bretsky, 1976, p. 273) compared with

about 1.0 for Cardiolucina species, and the shell is less tumid T/L

0.33 compared with 0.43 for Cardiolucina. Furthermore, R. amianta

has 8-12 broad radial ribs crossed by threadlike concentric lamel-

lae, often with intermediate secondary radial ribs in the ventral two

thirds of the shell. We agree with Keen (1971) and Britton (1972)

that Radiolucina amiantua belongs to a distinct and separate clade

which includes several fossil species from the south western USA.

Therefore, Bretsky’s (1976, p. 272) diagnosis of Bellucina based

on P. amiantus should be rejected.

Some American palaeontologists (Woodring, 1925; Gardner,

1926) used the generic name Cardiolucina for some Miocene luci-

nids with high beaks and lacking radial ribs. These species are now

placed in the genus Cavilinga Chavan, 1937 (see Olsson &

Harbison, 1953 p. 85; Bretsky, 1976 p. 265).

Chavan (1937, 1969) classified Bellucina as a subgenus of Linga

de Gregorio 1885, for which the type species is Lucina columbella

Lamarck, 1818, a Miocene fossil from France. This is a large, very

thick-shelled species with concentric ribs only and a strong posterior

sulcus unlike Cardiolucina.

SPECIES DESCRIPTIONS

Cardiolucina agassizi (Michelotti, 1839)

Fig. 1

Cardium agassizi Michelotti, 1839: 17.

Lucina agassizi — Michelotti, 1847: 404 pl. 4 figs 4,5, 7.

Lucina irregularis Eichwald, 1853: 82-83, pl. 5 fig. 4.

Cardiolucina agassizi (Michelotti) — Sacco, 1901: 89-90, pl. 10 figs

37-39.

Phacoides (Cardiolucina) agassizi (Michelotti) - Cossmann &

Peyrot, 1911: 688-689, pl. 28. figs 83-86.

TYPE MATERIAL. ?Museo di Paleontologia, Rome.

TYPELOCALITY. Not givenby Michelotti, but Sacco (1901) records

several localities in the Miocene of Italy.

DESCRIPTION. Shell small, solid, height to 7.4 mm, subcircular,

mean H/L 1.03, moderately inflated, mean T/L 0.39, tumidity to a

maximum of 2.7 mm on a single valve. Shell inequilateral, ex-

tended anteriorly. Deep posterior sulcus with fine concentric

lamellae. Shallow anterior sulcus. Lunule heart-shaped, small and

shallow. Escutcheon lanceolate with concentric lamellae and a

single strong radial ridge. Exterior sculpture of more than 25 faint,

radial, rounded ribs with narrow interspaces. There are about 20

prominent, concentric lamellae which are often slightly recurved.

Lamellae are extremely variable in thickness, degree of projection,

recurvature and width of interspaces. Hinge plate thick. Left valve

with single posterior lateral tooth, two cardinal teeth and single

anterior lateral. Right valve with single posterior and anterior

laterals and single cardinal. Ligament extending two thirds of the

distance from umbone to posterior lateral tooth. Inner margin with

28-30 crenulations and small denticles along anterodorsal and

posterodorsal margin. Anterior adductor muscle scar elongate, not

buttressed, with pallial line attached near to the ventral tip. Poste-

rior muscle scar ovate. Pallial line indistinct. Shell white.

In the material examined from France we found the concentric

ornament to be extremely variable from separated, thin lamellae to

clusters of thicker lamellae producing a rugose ornament (see Fig.

J.D. TAYLOR AND E.A. GLOVER

= *

Fig. 1 Cardiolucina agassizi (Michelotti, 1839), middle Miocene (Langhian), St Jean de Marsacq, Landes, France. (coll Lozouet and Maestrati, MNHN )

a, exterior of right valve; b, exterior of right valve; c, interior of left valve; d, interior of right valve. Scale bars a = 750um, b =700um, c = 1.5mm, d=

500um.

1d). Similar variability of the concentric sculpture is seen in the

Recent species C. crassilirata and C. semperiana.

DISTRIBUTION. As well as the records of Sacco (1901), C. agassizi

is also recorded from the Tortonian of the Aquitaine Basin (Coss-

mann & Peyrot, 1911) and from the middle Miocene Korytnica

Formation of Poland (Friedberg, 1934).

MATERIAL EXAMINED. FRANCE: Middle Miocene (Langhian),

St Jean de Marsacq, Landes, France; ca 100 shells, Lozouet &

Maestrati coll (MNHN).

Cardiolucina australopilula Taylor and Glover, new

species

Figs 2-6

TYPE MATERIAL. Holotype: WAM reg number 1 10—96: H. 13.6, L.

12.8, T. 6.6 mm. Paratypes: WAM 111-96 2 preserved specimens

station 35; AM C.202975 1 preserved specimen, station 23; BMNH

1996086, 1996087, 1996088, 1996089 4 preserved specimens, sta-

tions 1, 33, 36, 37; BMNH 1996090 one dry shell station 36. (Station

data in Glover & Taylor, 1997).

TYPE LOCALITY. 9 km north of Beacon Island, Wallabi Group,

Houtman Abrolhos Islands, Western Australia, 113° 46'E, 28° 23'S,

depth 39m.

SHELL DESCRIPTION. Shell solid, small, height to 13.8 mm,

subcircular in outline, H/L mean 1.03, inflated, orbicular, T/L mean

0.47, tumidity to a maximum of 7.5 mm on a single valve. Shell

slightly inequilateral, extended anteriorly. Very shallow posterior

sulcus. Umbones pronounced, prosogyrate. Lunule extremely small

and shallow. Anterodorsal area slightly convex, heart-shaped in

outline and with fine concentric ribbing. Escutcheon broadly lan-

ceolate, convex and finely ribbed with concentric and a few radial

ribs. Exterior sculpture of about 20 radial ribs, rounded in profile,

separated by flat interspaces. Anteriorly, interspaces are as wide as

the ribs, elsewhere interspaces are about two thirds of the width of

radial ribs. Radial ribs have very fine radial striations (Fig. 3d).

Radial ribs are crossed by 34—36 thin, low, concentric lamellae and

their intersection produces a finely fenestrate ornamentation. Hinge

plate thick, left valve with single posterior lateral tooth, two cardinal

teeth, the anterior of these is robust and hooked (Fig. 3c) and a single

anterior lateral. Right valve with single posterior lateral

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

Fig. 2 Cardiolucina australopilula Taylor and Glover new species, Beacon Island, Wallabi Group, Houtman Abrolhos Islands, Western Australia. a,

Holotype (WAM 110-96), exterior of right valve; b, Holotype, exterior of left valve; c, Paratype (BMNH 1996090), exterior of left valve; d, Paratype

(BMNH 1996090), interior of left valve; e. interior of right valve of juvenile shell, Eagle Bluff, Peron Point, Shark Bay; f, Holotype dorsal area. Scale

bars a—d = 2.0mm, e = 2.5mm, f = 2.0mm.

tooth, two cardinal teeth, the posterior larger than the anterior, and a

single anterolateral. Cardinal teeth sit on a curved, ventrally project-

ing buttress. Ligament elongate, extending from beak to posterior

lateral tooth. Inner margin of shell thickened with 21 crenulations.

Small sinus on posterolateral margin corresponding to the position

of the posterior sulcus. Anterior adductor muscle scar narrow,

elongate and sited on a buttress with ventral third detached from the

pallial line. Posterior muscle scar ovate. Pallial line continuous.

Shell colour white.

Juvenile shells (Fig 2e) less inflated, more anteriorly extended.

External radial ribs visible on interior of shell.

ANATOMY. The mantle margin is thick and fused only at the

posterior exhalant aperture (Fig. 4), and there is no fused inhalant

J.D. TAYLOR AND E.A. GLOVER

Fig.3 Cardiolucina australopilula new species. a, Holotype, hinge in the right valve; b. Holotype, hinge in the left valve; c, Paratype, left valve, detail

of cardinal teeth; d, outer shell surface showing radial striations on radial ribs. Scale bars a=2.0mm, b=2.5mm, c=1.0mm, d=500ym.

aperture. The exhalant aperture consists of a muscular tube with a

flared distal end, which is inverted in preserved specimens, but can

presumably be protracted. The ctenidia comprise inner demibranchs

only. These have large, thick filaments which are fused between the

ascending and descending lamellae. To the posterior, the outer

lamellae are attached to the mantle margin just ventral to the

exhalant aperture. The attachment area forms a thin, muscular

septum. The labial palps consist of small folds at the edge of the thin,

ridge-like lips. The foot is cylindrical and vermiform, tapering to a

blunt point and extending posteriorly with a rounded heel. The

visceral mass is covered by a muscular wall with a single, large,

hemispherical visceral lobe on each side. The rectum runs around

the dorsal side of the posterior adductor muscle and opens into the

mantle cavity close to the exhalant tube.

COMPARISON WITH OTHER SPECIES. This species is similar in size,

shape and sculpture to C. quadrata (Prashad, 1932) (Fig. 16) but

differs in the greater prominence of the radial ribs and thinner,

concentric lamellae. C. australopilula is also more tumid and spheri-

cal. This species has been referred to as Bellucina cf. semperiana by

Slack-Smith (1990), but semperiana 1s smaller, with different orna-

mentation and hinge teeth.

HABITAT. Fine, well-sorted, calcareous sand between 30—40m, in

association with other bivalves Amusium balloti, Mimachlamys

australis and Rastafaria thiophila (Glover & Taylor 1997).

DISTRIBUTION. Western Australia, Houtman Abrolhos Islands to

Broome (Fig. 6).

ETYMOLOGY.

MATERIAL EXAMINED. WESTERN AUSTRALIA: Broome (AM

C.309449); Roebuck Bay, Broome (BMNH 1892.1.29.121);

Montebello Islands, Bunsen Channel, N. end of Trimoville I. 20°

23.8'S; 115° 32.43'E 3m (WAM); Barrow Island 20° 53'S, 115° 20'E

(WAM); Exmouth Gulf (WAM 3461-3468); Mauds Landing, N.W.

Cape, 3m (WAM); Turtle Beach, off N.W. Cape (AM C.309456);

Carnarvon (AM C.309455); Dirk Hartog Island, 10-20m (WAM

3475.68); Bernier Islands 24° 53.3'S 113° 17.5'E. (WAM); Shark

Bay, Peron Peninsula (WAM 249.94); Shark Bay, 5 km E.S.E. Cape

Heirisson, 18m (WAM); Shark Bay, Freycinet Estuary (WAM 252—

94). Houtman Abrolhos Islands: N. of Wallabi Islands, 40m (BMNH

acc 2384); Easter Group, 50m (WAM N-2777); Zeewyk Channel,

40m (WAM N-2822); Little North Island, Easter Group, 40m

(WAM); Rat Island, 30m (WAM wet material); East of Good Friday

Bay, 28° 39.5'S — 113° 49.5'E, 40m (WAM).

australopilula meaning “Australian little ball’.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

# 1

i hi ai

3mm

Fig. 4 Cardiolucina australopilula anatomy. Right mantle and ctenidium

removed. Abbreviations aa — anterior adductor muscle, cme — cut mantle

edge, exa — exhalant aperture, ext — exhalant tube, fet — fused ctenidium

to mantle, ft — foot, Ict — left ctenidium, lp — labial palps, me — mantle

edge, pa — posterior adductor muscle, ppr — posterior pedal retractor

muscle, r — rectum, vl — visceral lobe.

Fig.5 Semidiagrammatic transverse section through the posterior

exhalant area of C. australopilula showing the inverted exhalant tube

and the connection of the ctenidium to the mantle. Abbreviations: cm —

connection between mantle and ctenidium, ct — ctenidium, ea — exhalant

aperture, iet — inverted exhalant tube, Im — longitudinal muscle, me —

mantle edge, n — nerve.

Fig. 6 Geographical distribution of Cardiolucina australopilula new

species.

Cardiolucina civica (Yokoyama, 1927)

Figs 7-9

Cardium civica Yokoyama, 1927: 179. pl. 48, figs 3 & 4.

Dentilucina (Bellucina) hedleyi Prashad, 1932: 163-164, pl. 5, figs

17 & 18.

Bellucina civica (Yokoyama, 1927) — Kuroda, Habe & Oyama,

1971: 394, pl. 118 figs 17 & 18.

Lucina (Codakia) semperiana (Issel, 1869) — Melvill & Sykes,

1898: pl. 3, fig.1.

TYPE MATERIAL. C. civica syntypes, University Museum, Tokyo,

CM 24913-24914, see Makiyama (1959); C. hedleyi holotype, I

individual 2 valves (ZMA).

TYPE LOCALTIES. C. civica was described as a Pliocene fossil from

Nagaya and Kakuma in Kaga Province, Japan. C. hedleyi from the

Philippines, Sulu Archipelago, Siboga station 105 (6° 8'N, 121°

19'E), depth 275 m.

NOMENCLATURE. We have examined the holotype of C. hedleyi

from Indonesia, which is very similar to both Recent specimens of

C. civica from Japan and the fossils figured by Yokoyama (1927) and

consider them conspecific.

SHELL DESCRIPTION. Shell solid, small, height to a maximum of

10.6 mm; subcircular, mean H/L 1.02, inflated, mean T/L 0.40,

tumidity to a maximum of 4.5 mm on a single valve. Shell slightly

extended anteriorly. There is a shallow posterior sulcus which has

concentric ribs. Lunule small and shallow. Anterodorsal area weakly

defined. Escutcheon slightly convex and finely ribbed. Exterior

sculpture of numerous (more than 30) fine, slightly rounded, radial

ribs with narrow interspaces. Radial ribs are crossed by many (more

than 25) conspicuous concentric lamellae, which are rounded in

profile and slightly recurved. Left valve with a single posterior

lateral, two cardinal teeth and a single posterior lateral; right valve

with a single posterior lateral, two cardinal teeth, the posterior

J.D. TAYLOR AND E.A. GLOVER

Fig.7 a-d Cardiolucina civica (Yokoyama, 1927) southern Honshu, Japan BMNH 1878.10.16.191 and e-f Dentilucina (Bellucina) hedleyi Prashad,

1932 Holotype, Sulu Archipelago, Philippines. a, exterior of left valve; b, interior of left valve; c, hinge in left valve; d, outer surface of shell showing

concentric and radial ribs; e, Holotype C. hedleyi exterior of left valve; f, Holotype C. hedleyi interior of left valve. Scale bars a=2.5mm, b=1.2mm,

c=500um, d=500um, e=2.0mm, f=2.5mm.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

101

Fig. 8 Cardiolucina civica from Andaman Islands. Large specimen illustrated by Melvill and Sykes (1897, fig.1) as Lucina (Codakia) semperiana Issel

(BMNH 1898.4.30.7) a, Right valve exterior; b, Left valve interior. Scale bar a—b = 1.0mm.

cardinal large and the anterior very small. There is a single anterior

lateral tooth. Ligament short, extending from the beak to about half

way to posterior lateral tooth. Inner margin of shell thickened and

with 30 crenulations. Small denticles on anterodorsal margin. Ante-

rior adductor muscle scar elongate and not buttressed, pallial line

attached near ventral tip. Posterior adductor scar small and rounded.

Pallial line continuous. Shell grey white.

Juveniles less inflated and more extended anteriorly. Posterior

sulcus more pronounced and anterodorsal shell margin markedly

convex. Shell thinner with radial riblets visible from interior of shell.

A large specimen (BMNH 1898.4.30.7) from the Andaman

Islands, figured by Melvill and Sykes (1898 pl.3, fig.1) as Lucina

(Codakia) semperiana, has many of the characters of C. civica,

although larger than specimens from other localities. Other speci-

mens in the BMNH from Port Blair, Andamans are similarly

large.

Fig.9 Geographical distribution of Cardiolucina civica (Yokoyama).

102

J.D. TAYLOR AND E.A. GLOVER

Fig. 10 Cardiolucina crassilirata (Tate, 1886), Western Port, Victoria, Australia. BMNH 1963200. a, exterior of left valve; b, interior of left valve. Scale

bars a = 2.0mm, b = 2.5mm.

COMPARISON WITH OTHER SPECIES. C. civica is most similar to C.

crassilirata, but it has a shorter ligament and coarser marginal

crenulations.

HABITAT. Fine sandy and shelly bottoms mainly from shallow

water to 200m, with a few dead shells recorded up to 600 m.

DISTRIBUTION. Andaman Islands, North Western Australia, South

East Asia, Japan, to Fiji (Fig. 9).

MATERIAL EXAMINED. ANDAMAN ISLANDS: 11°37'N,

92°56’E, 357m (ANSP 292239); Port Blair, BMNH 1953.1.30.199—

201; BMNH 1898.4.30.7. AUSTRALIA: Western Australia, N.W.

of Port Hedland 19°13'S, 116°6'E, 271m & 19°24'S, 115°52'E,

238m (AM C.309458 & C.309453); N.W. Beagle Island, 29°43'S,

114°17'E, 274—283m (AM C.309465); N.W. Roebuck Bay, Broome

17°34'S 120°22'E, 188m (AM C.309457). PHILIPPINES: Luzon,

off Matocot Pt W., 370m (USNM 295697); Tayabas Bay, off Tay-

abas Lt, 200m (USNM 297745); Ragay Gulf, off North Burias,

185m (USNM 246286); off Balanja Pt, S.E. Mindoro, 280m (USNM

295581); Adyagan I. E. Masbate,190m (USNM 298134); off Pt

Dumurug, Masbate, 273m (USNM 298227); Linapacan Strait, 90m

(USNM 301994); off Pt Lauis, E. Cebu, 280m (USNM 300493); off

Gigantangan I., N.W. Leyte, 210m (USNM 298317); off Dammi I.,

Sulu Archipelago, 446m (USNM 299036); off Simaluc Island, Tawi

Tawi Is, 600m (USNM 299195); off Jolo, Jolo I., 283m (USNM

294618) & 37m (USNM 294584); 14°O1'N 120°18'E, 192m

(MNHN). CHINA: off Pratas Islands., China Sea, 272m, (USNM

296724). JAPAN: off southern Honshu 34°13'N, 136°13'E, 90—

100m (BMNH 1878.10.16.191); Kumano-Nada, off Kii Peninsula,

93m (NSMT 70534); off Jyogashima I., Miura Peninsula (NSMT

70535); off Goto Islands, 168m (NSMT 56297); off Tsushima

Islands, 112m (NSMT 56298); Usibuka, Anakusa Island (NSMT

70533); Sagami Bay (USNM 708071). IRIAN JAYA: Aoeri I.,

Geelvink Bay, 140 (ANSP 275808, 280332); Japen I., Geelvink

Bay, 30-40m (ANSP 277817, 276638, 279725); Schouten I., 18m

(ANSP 280204). NEW CALEDONIA: 19°07'S 163°22'E, 110-

200m (MNHN); 22°02'S 165°57'E 135-150m (MNHN); Grande

Récif Sud, 22°42'S 157°22'E, 35m (MNHN). FIJI: Viti Levu Bay 0—

5m near shore (USNM 878321).

Cardiolucina crassilirata (Tate, 1887)

Figs 10-11

Lucina crassilirata Tate, 1887a: 6—7, pl. 4 fig. 2.

Codokia crassilirata (Tate) — Cotton & Godfrey, 1938: 206 fig. 218.

Loripes crassilirata (Tate) — Pritchard & Gatliff, 1903: 139.

Bellucina crassilirata (Tate) - Macpherson & Gabriel, 1962: 327.

Fig. 11

Geographical distribution of Cardiolucina crassilirata (Tate).

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

Linga (Bellucina) crassilirata (Yate, 1887) — Lamprell & White-

head, 1992: pl. 21 figs 128a & b.

HOLOTYPE. South Australian Museum No. D.12957. L = 8.5 mm,

H = 8.3 mm, T = 7.0 mm.

TYPE LOCALITY. Streaky Bay, near Ceduna, South Australia.

SHELL DESCRIPTION. Shell small, solid, height to 8.4 mm, outline

circular, H/L mean 0.98; shell moderately inflated with T/L mean

0.41, tumidity of a single valve to a maximum of 3.4 mm. Shell

slightly inequilateral, extended anteriorly. Shallow sulcus poster-

iorly, with concentric lamellae. Lunule small and shallow. Escutcheon

slightly convex, with fine concentric ribs. Exterior sculpture of more

than 30 fine concentric lamellae, clustered in groups of 4—6 with

broader interspaces between the clusters. Radial sculpture of many

fine ribs visible in the interspaces of the concentric ribs. Hinge plate

narrow; left valve with single posterior lateral, two cardinal teeth and

single anterior lateral. Right valve with a single posterior and

anterior laterals and two cardinal teeth, posterior cardinal larger than

anterior cardinal tooth. Ligament long, extending from beaks to

posterior lateral tooth. Inner margin with around 60 fine crenulations.

Anterior adductor muscle scar, elongate, not buttressed, with ventral

quarter detached from pallial line. Posterior adductor muscle scar

ovate and not buttressed. Pallial line discontinuous. Shell greyish

white.

COMPARISON WITH OTHER SPECIES. See C. civica.

HABITAT. Shallow subtidal to 70m.

DISTRIBUTION. South Australia, Victoria and Tasmania (Fig. 11).

MATERIAL EXAMINED. AUSTRALIA: Victoria, BMNH

1906.9.19.70-2; Western Port (BMNH 1963200); BMNH

1911.12.19.11-12; Western Port (WAM); Western Port (AM

C.303440); Western Port, 15—18m (AM C.309469); Port Fairy,

beach (ANSP 135488); New South Wales, Between Eagle and

Crawfish Rock (AM); South Australia, Spencers Gulf (BMNH

1893.12.12.352-54); Hardwicke Bay (AM C.13463); Neptune L.,

70m (AM); South Australia (AM C.6677); Tasmania, Geography

Strait, 27m (AM C.38180).

Cardiolucina eucosmia (Dall, 1901)

Figs 12-13

Lucina pisum Reeve, 1850 (in part): pl. 11 fig. 66a (non Lucina

pisum Sowerby, 1836: 341).

Phacoides (Bellucina) eucosmia Dall, 1901: 806 (replacement name

for L. pisum Reeve).

Phacoides eucosmia Dall. — Hedley, 1909: 426, pl. 37 fig 16.

TYPE MATERIAL. Lectotype here selected BMNH 1963194/1 (Figs

12a—d). Paralectotypes BMNH 1963194/2-3; 1845.825.210-212;

1963552 (this is C. semperiana see p. 107)

TYPE LOCALITY. Restricted to Port Essington, Northern Territory,

Australia (modified from Reeve, 1850).

NOMENCLATURE. Many authors, for example Fischer (1871), Lamy

(1920), Chavan (1969), Britton (1972), Bretsky (1976) and Oliver

(1992) have considered Lucina semperiana Issel, 1869 a senior

synonym of P. eucosmia. However, Lucina pisum Reeve, for which

Dall gave the replacement name Phacoides eucosmia, is based on a

syntype series composed of two distinct species. For the reasons

given above in the generic section, we consider the specimens from

Port Essington to conform most closely with Reeve’s description

103

and concept of Lucina pisum and one of these has been selected as a

lectotype. The other syntypes from Singapore, now become

paralectotypes and we consider these to be the eastern Indian Ocean

form of C. semperiana (Fig. 21 a & b).

SHELL DESCRIPTION. Shell solid, small, height to 7.8 mm,

subcircular in outline, slightly extended in an anterolateral direction,

H/L mean 1.05; inflated, T/L mean 0.49, tumidity to a maximum of

3.8 mm on a single valve. Posteriorly, there is deep sulcus extending

from hinge to ventral margin. The sulcus and the posterior portion of

the shell has fine concentric ribbing with no radial ribs. The lunule

is ‘U-shaped and deeply incised to the ventral edge of the hinge

plate. Anterodorsal area is slightly convex, ovate in outline with fine

concentric sculpture. Posterior dorsal area broadly lanceolate in

outline, convex, with fine concentric ribbing. Exterior sculpture of

about 12 broad, radial ribs with rounded profile, somewhat flattened

on upper surface. Ribs narrower towards umbos. Interspaces between

ribs deep, narrower than ribs. Radial ribs crossed by 13-14 thin,

low, concentric lamellae which project slightly above radial ribs.

The intersection of concentric and radial sculpture gives a distinctive

fenestrate ornament. Hinge plate thick. Left valve has a single,

strong lateral tooth posteriorly, two thin, cardinal teeth, the posterior

tooth much larger than the anterior. Cardinal teeth are denticulate

(see Fig. 12d). A single anterior lateral lies close to perimeter of the

lunule. Right valve has a single posterior lateral, a large single

cardinal tooth and a single anterior lateral. Both valves have small

denticles around the dorsal margin of valve. Ligament elongate,

extending from beak to posterior lateral tooth. Inner shell margin

thickened, with 12 coarse crenulations. Small sinus on posterior

lateral margin corresponding to position of sulcus. Anterior adduc-

tor muscle scar elongate, sitting on a shallow buttress with the only

the ventral tip detached from the pallial line. Posterior adductor scar

ovate on a shallow buttress. Pallial line is thin and continuous. Shell

colour creamy white.

Juvenile shells (Figs 12 e & f) are less inflated and more extended

anteriorly. The lunule is widely opened in younger stages. Denticles

are absent on the dorsal inner margins. Radial ribs begin narrow and

broaden ventrally between each concentric rib. Radial ribs are

visible on interior of shells.

COMPARISON WITH OTHER SPECIES. Although C. pisiformis, C.

siquijorensis and C. australopilula have a similar fenestrate orna-

ment, C. eucosmia can be easily distinguished from all other species

by the deeply incised lunule.

HABITAT. Low intertidal to 100 m in fine sand and mud. Hedley

(1909) reports dredging the species in the Gulf of Carpentaria from

about 10 m in soft black mud, where it was abundant.

DISTRIBUTION. Northern Australia (Fig. 13).

MATERIAL EXAMINED. AUSTRALIA: Northern Territory and

Queensland, BMNH 1963490, 1963236, 1910.9.28.77-8; W. of

Cape York, 8m (BMNH 1887.2.9.2776-7); Gulf of Carpentaria

(NMW 1955.158); Sweers I. (AM C.75249, 78223); Forsyth I. (AM

C.14892); Horsey River mouth, 9m (AM C.15152), Karumba (AM

C.14963); Mapoon (AM C.14281); off Bountiful Islands (AM);

Gulf of Carpentaria (AM C.100498); mid-Gulf of Carpentaria, 66m

(AM); off Albany River (AM); Karumba Point (AM 1909); Albany

Passage, 8-25m (AM C. 36172); Port Essington (BMNH,; off Point

Charles, Darwin, 27-37 m (AM C.309468); Van Diemen’s Inlet

(NMW 1955.158); Van Diemen’s Inlet (AM C.15273); 100 miles N.

of Croker Island, Arafura Sea, 124m (AM C.309454): 150 km N. of

Coburg Peninsula, 108m (AM); Dinah Beach, Darwin (AM); Cro-

codile Research Station, Manningrida, Arnhem Land (AM);

J.D. TAYLOR AND E.A. GLOVER

Fig. 12 Cardiolucina eucosmia (Dall, 1901) Port Essington, Northern Territory, Australia. Figured specimens a—d are the lectotype of Lucina pisum

Reeve, BMNH 1963194/1. & Figure e-f C. eucosmia, Arafura Sea, Northern Territory, Australia, AM C.309454. a, exterior of right valve; b, exterior of

left valve; c, interior of left valve; d, hinge and lunule in left valve; e, juvenile, interior of left valve; f, juvenile, exterior of left valve. Scale bars a—c =

1.0mm, d = 500um, e = 650um, f = 1.0mm.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

Fig. 13 Geographical distribution of Cardiolucina eucosmia (Dall).

Western Australia, Prince Frederick Harbour (WAM); Cape Leveque

(AM); Port Hedland (AM); Direction Island, off Onslow (AM

C.69334).

Cardiolucina pisiformis (Thiele, 1930)

Figs 14-15

Phacoides (Parvilucina) pisiformis Thiele, 1930: 592.

Phacoides (Parvilucina) pisiformis Thiele — Ponder, 1978: 439, pl. 2

figs 3 & 9.

Bellucina pisiformis (Thiele) — Slack-Smith, 1990: 135.

HOLOTYPE. Museum fiir Naturkunde, Berlin. Reg. no. 67726. L=

3 mm, H= 2.9 mm (details from Ponder, 1978).

TYPE LOCALITY. 5 km N.W. of Denham, Shark Bay, Western

Australia, 3 m.

SHELL DESCRIPTION. Shell very small, toa maximum height of 4.4

mm, outline circular, H/L mean 0.97; inflated, T/L mean 0.42,

tumidity of single valve to 2.0 mm. Inequilateral, slightly extended

anteriorly. Shallow posterior sulcus with concentric lamellae. Lunule

heart-shaped, slightly excavated, smooth. Anterodorsal area indis-

tinct. Posterodorsal area with concentric lamellae. Exterior sculpture

of 13-17 low, flattened, radial ribs with broad interspaces. Ribs

broader and more rounded towards anterior of shell. About 20 thin,

concentric lamellae project slightly above radial ribs. Sculpture

distinctly fenestrate in appearance. Hinge plate strong. Left valve

with single posterior lateral, two cardinal teeth, the posterior of these

large and robust, the anterior small. Single anterior lateral tooth.

Right valve with a single, strong, posterior lateral tooth, two cardi-

nals, the anterior large and the posterior very small and a single

anterior lateral. Ligament extends from beak to posterior lateral

tooth. Inner margin with about 13 coarse crenulations, which are

sometimes bifurcate with occasional small denticles in the

interspaces. Small, rounded denticles around anterior and posterior

dorsal margins. Anterior adductor muscle scar elongate, slightly

buttressed with only the ventral tip detached from the pallial line.

Posterior adductor muscle scar rounded. Pallial line continuous.

Shell white.

Juvenile shells (Fig.14 e) are more extended both anteriorly and

posteriorly, with a prominent convex anterodorsal area.

105

COMPARISON WITH OTHER SPECIES. This is a small distinctive

species with an exterior ornament similar to C. eucosmia and C.

siquijorensis. It is smaller, more circular in shape and lacks the

deeply-excavated lunule of C. eucosmia. C. siquijorensis has fewer

and broader radial ribs, a shallower lunule and a different dentition.

HABITAT. Shallow water to 240 m.

DISTRIBUTION. Western Australia (Fig. 15).

MATERIAL EXAMINED. AUSTRALIA: Western Australia, W. of

Rottnest Island, 110m (WAM 262-94); Point Peron Peninsula,

Shark Bay (WAM 257-94, 258-94, 259-94, 260-94,); Carnarvon

(WAM); N. of Homestead Point, Dirk Hartog Island (WAM 250-

94); Ningaloo Reef, N.W. Cape (WAM); Mangrove Bay beach,

N.W. Cape (WAM); Yardie Creek to Tantibiddi, N.W. Cape (WAM

wet material); N.W. of Port Hedland, 112m (AM C.309462):; N.W.

Shelf, W. of Roebuck Bay, 238m (AM C.309464); Broome (ANSP

233737); Northern Territory, N. of Cobourg Peninsula, 55m

(AM); 200km N. of Melville Island, 215m (AM).

Cardiolucina quadrata (Prashad, 1932)

Figs 16-17

Dentilucina (Bellucina) hedleyi var. quadrata Prashad, 1932: 164,

pl. 5 figs 19 & 20.

TYPE MATERIAL. Holotype, one individual two valves, L=10.5

mm, H=10.5 mm. Also 4 paratype valves. (ZMA).

TYPE LOCALITY. Siboga Expedition station 212 (5° 54.5'S, 120°

19.2'E), west of Salayar, Sulawesi, Indonesia, 462 m.

SHELL DESCRIPTION. Shell solid, small, height to 12.9 mm,

subcircular, H/L mean 0.97, moderately inflated, mean T/L 0.38,

tumidity to a maximum of 5.3 mm on a single valve. Shell

inequilateral, extended anteriorly. Shallow posterior sulcus with

concentric lamellae; umbones prominent. Lunule extremely small

and shallow. Anterodorsal area heart-shaped, slightly convex with

fine concentric lamellae. Escutcheon lanceolate, with fine concen-

tric lamellae and a few radial ribs. Exterior sculpture of about 25

fine, closely spaced, radial ribs which are rounded in profile. Radial

ribs crossed by 25-30 low, broad, concentric lamellae, the intersec-

tion giving a finely beaded appearance. Hinge plate thick. Left valve

with an single posterior lateral tooth, two cardinal teeth, the anterior

of which is hooked and sits on a rounded buttress, and a single

anterior lateral tooth. Right valve with single posterior lateral, two

cardinal teeth with the posterior larger and a single anterior lateral

tooth. Ligament elongate extending from beak to two thirds of

distance to the posterior lateral tooth. Inner margin with about 25

crenulations, some of those to the posterior bifurcating. Shallow

sinus on posterior margin. Anterior adductor muscle scar elongate,

not buttressed, slightly detached at the ventral tip from the pallial

line. Posterior adductor muscle scar ovate. Pallial line continuous.

Shell colour white.

COMPARISON WITH OTHER SPECIES. Although this species name

was first introduced by Prashad (1932) as a variety of Dentilucina

(Bellucina) hedleyi = C. civica) it is clearly distinct. In many

characters, such as hinge teeth, it resembles C. australopilula, but

differs in shape and external sculpture, the latter having a fenes-

trate ornament compared with the finely beaded sculpture of C.

quadrata.

HABITAT. A deeper water species from muds at 230-1280 metres.

J.D. TAYLOR AND E.A. GLOVER

Fig. 14 a—d, f Cardiolucina pisiformis (Thiele, 1930) Bandicoot Bay, Barrow Island, Western Australia, WAM 268-94 and Fig. e C. pisiformis, Peron

Peninsula, Shark Bay, Western Australia WAM 258-94. a, exterior of right valve; b, exterior of left valve; c, interior of left valve; d, interior of right

valve; e, juvenile, exterior of shell; f, exterior of shell surface showing posterior sulcus. Scale bars a—d = 1.0mm, e = 500um, f = 300um.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

Fig. 15 Geographical distribution of Cardiolucina pisiformis (Thiele).

DISTRIBUTION. Indonesia and Philippines (Fig. 17).

MATERIAL EXAMINED. INDONESIA: Kai Islands, 05°23'S

132°37'E, 436-413m & other stations in same area between 250-—

450m (MNHN, Bouchet and von Cosel manuscript).

KALIMANTAN: Sibuko Bay, 762 & 589m (USMN 302240).

PHILIPPINES: 11°58'N 122°02'E, 486-55 1 m (MNHN) (see Cosel

& Bouchet, ms.); off Baliscasag I.,S. Bohol, 812m (USNM 298742);

Tligan Bay, N. Mindanao, 500m (USNM 298616) & 505m (USNM

298660): off Claveria, Burias, 390m (USNM 2800546): off N. Ticao

1.,420m (USNM 298025); off Ponson I., Camotes Islands, 650m

(USNM 298468); off Apo I., S. Negros, 460m (USNM 298759);

Sogod Bay, Leyte,1280m (USNM 298560).

Cardiolucina rugosa (Hedley, 1909)

Figs 18-19

Lucina (Myrtea) seminula Gould, 1862: 174 (non Lucina seminulum

Deshayes, 1858: 673).

Lucina (Codakia) seminula (Gould) — Smith, 1885: 180-181, pl. 13,

figs 5 & Sa.

Phacoides rugosus Hedley, 1909: 427.

Lucina seminula Gould — Johnson, 1964: 148, pl. 28 fig. 3.

TYPE MATERIAL. Lectotype selected by Johnson (1964, pl 28 fig.

3) (USNM 553). Hedley (1909) introduced the replacement name P.

rugosus because L. seminula is preoccupied. The specimen from the

type locality illustrated in Fig. 18 was given by Gould to Cuming.

TYPE LOCALITY. Hong Kong Harbour, 9-18 m.

SHELL DESCRIPTION. Shell solid, small, height to a maximum of

3.1mm, subcircular in outline, H/L mean 0.92; moderately inflated,

T/L mean 0.38, tumidity to a maximum of 1.2 mm ona single valve.

Shell inequilateral and extended towards anterior. Pronounced pos-

terior sulcus with concentric lamellae. Anteriorly, there is a narrow

sulcus. Lunule small, heart-shaped. Anterodorsal area shallowly

concave with fine concentric lamellae. Escutcheon lanceolate and

slightly convex. Exterior sculpture of about 20 thick, slightly

recurved, concentric lamellae. Interspaces between concentric la-

107

mellae are variable in width. Concentric lamellae often extended

into short flanges at margin of posterior sulcus. Radial ribs indis-

tinct, flattened and closely spaced. Hinge plate thick. Left valve with

single posterior lateral, two cardinal teeth, the anterior of which is

thicker and triangular and a single anterior lateral. Right valve with

a single posterior lateral tooth, two cardinal teeth, the posterior large

and the anterior a thin plate, with a single anterior lateral tooth.

Ligament very short, extending from beak to less than half way to

posterior lateral tooth. Inner margin with 20-21 sharp crenulations.

Small denticles on posterior and anterior dorsal margin. Shallow

sinus on posterior margin corresponding to the position of the

posterior sulcus. Anterior adductor muscle scar elongate, not but-

tressed, detached from the pallial line only at the ventral tip. Posterior

adductor muscle scar ovate. Pallial line continuous. Shell colour

creamy white.

COMPARISON WITH OTHER SPECIES. Cardiolucina rugosa is closely

similar to C. semperiana, but it is smaller, less tumid with more

prominent, recurved concentric lamellae and greater anterior exten-

sion.

HABITAT. 10—250 m witha few dead shells from 540 m, sublittoral

silts and muds.

DISTRIBUTION.

MATERIAL EXAMINED. HONG KONG: North Lantau (BMNH;

BMNH 1963233; NMW acc. 55.158; AM C.33806; 9-18m (USNM

611842, 2 paratypes); (USNM 24252); (USNM 714821). CHINA:

off Pratas Islands, 268m (USNM 296955). KALIMANTAN: off

Silungan I., 539m (USNM 291027). PHILIPPINES: (USNM

21109); Pescador I. Tanon Straits, 412m (USNM 293876): Bohol

(BMNH); off Sombrero I., Balayan, 216m (USNM 312342); S.E. of

Bantayan I., 60m (USNM 293115); Destacado I., 218m (USNM

298090); off N. Cebu, 125m(USNM 293266); Pujada R., E.

Mindanao, 253m (USNM 294765); off Tochanhi Pt, Tawi Tawi

Islands, 90m (USNM 292997) & 35m (USNM 283372); 11°43'N,

122°34'E, 93-99m (MNHN). AUSTRALIA: Queensland, Cape

York (BMNH 1887.2.9.2772-5); Hope Islands, 10-18m (BMNH

1910.9.28.68-71 and BMNH 1963259); Palm Island (NMW acc.

55.158); Gulf of Carpentaria 16°58'S 140°53'E, 9m (AM C.015275);

Albany Passage, Torres Strait, 9-30m (AM C.36171); Palm Island

(AM C.1049); Low Isles, 27m (AM); Cape Sidmouth (AM C.

2594); Endeavour Reef, 40m (AM C.44668); Hervey Bay (AM);

Barney Point, Port Curtis (AM C.21805); Facing Island (AM C.4428);

Quoin Island, Port Curtis (AM); Northern Territory, Darwin (AM).

PAPUA NEW GUINEA: 0.5 mile off Fairfax Harbour, Port Mores-

by, 15—18m (AM C.309466).

Queensland, Australia to Hong Kong (Fig. 19).

Cardiolucina semperiana (Issel, 1869)

Figs 21-24

Lucina pisum Reeve 1850 (part), pl. 11, fig. 66b (non L. pisum

Sowerby 1836).

Lucina semperiana Issel, 1869: 82-83.

Phacoides (Bellucina) semperiana (Issel) — Lamy 1920: 211-213.

Dentilucina (Bellucina) macassari Prashad 1932: 163 pl. 5 figs 13-

16.

Lucina semperiana (Issel) — Bouchet & Danrigal 1982: 16, fig. 19.

Bellucina semperiana (Issel) — Oliver 1992: 99, pl. 19 fig. 7.

TYPE MATERIAL. ‘The species concept of C. semperiana was based

on the figure in Savigny (1817, pl. 8, fig 12) and one specimen from

Suez in the Issel Collection (now lost); the single shell of this species

in the Savigny collection (MNHN) illustrated by Bouchet & Danrigal

J.D. TAYLOR AND E.A. GLOVER

Fig. 16 Cardiolucina quadrata (Prashad, 1932). a—d, holotype of Dentilucina (Bellucina) var. quadrata Prashad, 1932, Salayar, Sulawesi, Indonesia and

e-f, C. quadrata, Kai Islands, Indonesia, BMNH. a, exterior of right valve; b, exterior of left valve; c, interior of left valve; d, interior of right valve; e,

exterior shell surface; f, hinge showing hooked anterior cardinal tooth in the left valve. Scale bars a-e = 2.0mm, f = 1.0mm.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

Fig. 17 Geographical distribution of Cardiolucina quadrata (Prashad).

(1982, fig. 19) is here designated as lectotype. The holotype of

Dentilucina (Bellucina) macassari Prashad is located at ZMA, reg.

number 3.32.090 (Fig. 21 c & d).

TYPELOCALITY. C. semperiana from Suez, Red Sea. C. macassari

from Makassar, South Sulawesi, Indonesia (Siboga Expediton sta-

tion 71).

SHELL DESCRIPTION. Shell solid, small, height to a maximum of

7.7 mm, sub-circular in outline H/L mean 0.98; inflated, T/L mean

0.46, tumidity of single valve to 4.6 mm. Inequilateral, extended

109

Fig. 19 Geographical distribution of Cardiolucina rugosa (Hedley).

anteriorly. Deep sulcus posteriorly with prominent concentric la-

mellae. Posterodorsal margin of the sulcus marked by a ridge with

prominent short lamellae. Lunule heart-shaped, shallow. Anteriorly,

there is a narrow sulcus. Escutcheon lanceolate with fine concentric

sculpture. Exterior sculpture of 20—30 low, rounded radial ribs with

narrow interspaces. These are crossed by 15—24 projecting, concen-

tric lamellae which are often irregularly spaced. In some specimens,

concentric lamellae are thickened with steep, ventral edges, giving a

stepped appearance to the shell. Hinge plate thick. Left valve with

single posterior and anterior lateral teeth and two cardinal teeth, of

Fig. 18 Cardiolucina rugosa (Hedley, 1909), Hong Kong Harbour, BMNH 1963233. Specimen of Lucina seminula Gould 1862 from type locality, given

by Gould to Cuming. a, exterior of left valve; b, interior of right valve. Scale bar a—b = 1.0mm.

J.D. TAYLOR AND E.A. GLOVER

Fig. 20 a—b Cardiolucina semperiana (Issel, 1869), Gulf of Suez, Red Sea BMNH 1870.1.19.4, c-f C. semperiana, Tuléar, Madagascar, BMNH. a,

exterior of right valve; b, interior of left valve; c, dorsal view articulated shell; d, exterior of right valve; e, interior of left valve; f, hinge showing details

of the dentition with denticulate posterior cardinal tooth. Scale bars a—b = 1.0mm, c = 750um, d—-e = 1.0mm, f = 500um.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

111

Fig. 21

Cardiolucina semperiana (Issel, 1869). Figure a—b Paralectotype of Lucina pisum Reeve, 1850 BMNH 1963552, Singapore and c—d Holotype of

Dentilucina (Bellucina) macassari Prashad, 1932, Makassar, Indonesia, ZMA. a, exterior of right valve; b, interior of left valve; c, exterior of right valve;

d, interior of left valve with attempted drill hole on the posterior part of the umbo. Scale bars a—d = 1.0mm.

which the anterior is the smaller. Right valve with single posterior

and anterior laterals and two cardinals, the posterior being the larger.

Cardinal teeth have small denticles. Ligament extends two thirds of

the distance from the beak to the posterior lateral tooth. Inner margin

with 20-21 crenulations and there are also small, rounded denticles

along anterodorsal and posterodorsal margins. Both anterior and

posterior shell margins have shallow sinuses corresponding to the

sulci. Anterior adductor muscle scar elongate, buttressed in the

more tumid specimens, detached from the pallial line only at the

ventral tip. Small pedal retractor muscle scar at dorsal tip of anterior

adductor scar. Posterior adductor muscle scar ovate. Pallial line

continuous. Shell creamy white.

ANATOMY. (Fig. 22 ) Posterior inhalant aperture absent, exhalant

aperture a small inverted tube. Ctenidia large and thick, inner

demibranchs only, with a thin, shallow, ventral food groove. Gill

filaments long and broad, with large central space crossed by

transverse muscle strands (Fig. 22 c). Bacteriocyte zone packed with

ovoid bacteria about 2.0—2.5 um in length (Fig. 22 d). Labial palps

consist of small papilla at edge of lips. Foot cylindrical, vermiform,

with a much-corrugated outer surface, a pointed tip and a small

rounded heel to the posterior. Ctenidia attached to posterior mantle

by a thin muscular sheet (Fig. 22 f). Body wall with a large irregular

visceral pouch in the antero-ventral position.

GEOGRAPHICAL VARIATION. This species is extremely variable

both within populations and across its range from the northern end of

the Red Sea to the Philippines. In Southeast Asia semperiana are

generally smaller, but relatively more tumid, as shown in Fig. 23,

which plots ratio of length over tumidity against mean height of

specimens from Red Sea, Madagascar, India, South East Asia. The

holotype of macassari is the most tumid of all the specimens

measured, but is probably an abnormal individual which has sur-

vived attempted drilling at the posterior dorsal margin. C. semperiana

from Southeast Asia also tend to have fewer, but heavier, concentric

lamellae (Figs. 21).

HABITAT. Live material collected from shallow water (up to 50 m),

a few dead shells from up to 350 m, mud and sand.

COMPARISON WITH OTHER SPECIES. C. semperiana is similar to C.

rugosa but is larger, more tumid and has more prominent radial ribs.

112 J.D. TAYLOR AND E.A. GLOVER

Fig. 22 Cardiolucina semperiana (Issel, 1869), Sinai Coast, Gulf of Suez, Egypt. NUW 1982.068. a, interior of right valve showing thick inner

demibranch of the ctenidium and cylindrical, vermiform foot; b, right valve showing pronounced visceral pouch and the foot with strongly corrugated

surface and small, rounded heel; c, section showing broad gill filaments and transverse muscle strands crossing the spaces within the filaments.

d, bacteriocyte zone with ovoid bacteria; e, labial palps (Ip) and mouth (m); f, ctenidia attached to the posterior mantle by a thin muscular sheet.

Abbreviations: c=ctenidium, f=foot, v=visceral pouch, h=heel, lp=labial palps, m=mantle, a=attachment of ctenidium to mantle. Scale bars a = 1.0mm, b

= 500um, c = 40um, d = Sum, e = 150um, f = 250um.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA 113

8.0

7.5

7.0

6.5

e SEAsia

= 6.0 oO Arabia

@ 5.5 4 Madagascar

5.0 © India

Vv macassari

4.0

3.5

3.0

co, of 40 42 44 AG .48 £50 .52 54 56

Tumidity/length

Fig. 23. Geographical shape variation of Cardiolucina semperiana. The specimen of C. macassari is the holotype.

Fig. 24 Geographical distribution of Cardiolucina semperiana (Issel).

114

J.D. TAYLOR AND E.A. GLOVER

Fig. 25

Cardiolucina siquijorensis Taylor and Glover new species, Siquijor, Philippines. a, Paratype (USNM 812176), exterior of right valve; b, Holotype

(USNM 812176), exterior of left valve; c, Paratype, interior of left valve; d, Holotype, detail of lunule; e, Paratype, hinge of left valve showing posterior

lateral tooth with pronounced rim on the socket. Scale bars a—c = 500um, d = 150m, e = 250um.

DISTRIBUTION. Northern Red Sea, southern Africa to Philippines

(Fig. 24).

MATERIAL EXAMINED. EGYPT: Gulf of Suez (BMNH

1870.1.19.4); Suez (BMNH 1963492): Nuweiba, Sinai (AM 1954).

YEMEN: Aden (BMNH 1902.12.30.754-6 & 1963235); Aden

(AM C.33727); (USNM 608854). OMAN: Qurm, Muscat (NMW

Z.1993.061.0434); Masirah (NMW Z.1993.061.0427 & 0433).

ARABIAN GULF: (USNM 636963); Bahrain (NMW

1993.061.043); Linyah (NMW acc. 55.158); Qatar (BMNH acc.

2258); Abu Dhabi (BMNH). ZANZIBAR: Nyange I., 20m (ANSP

251213); 1.5 miles S.W. of Ras Shangani, 18—20m (ANSP 213493);

1 mile N. Bawi I., 22m (ANSP 250912). KENYA: Mombasa

(BMNH acc. 2223), (NMW 1955.158). TANZANIA: Msimbasi

Bay, Dar es Salaam 5m (AMC.309448). MOZAMBIQUE: Bazaruto

I., 8-10 m (NM K5317 & G4458). SOUTH AFRICA: Durban Bay,

(NM E834 & A2357); Shelly Beach, Izotsha, Natal (NM9898).

MADAGASCAR: N.W. Madagascar, 30m (AM C.70660); Tuléar

(MNHN); 40 miles W.S.W. Tuléar, 33m (USNM 761189); Nossi Bé

(USNM 633927); Nossi Bé, shallow water to 35m, (ANSP 261127,

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

259587, 259028, 260541, 261338, 259901, 261873). MAURI-

TIUS: 1.5m N.W. Black River, 15m (ANSP 273431). PAKISTAN:

Karachi (BMNH acc. 1831). INDIA: 80 miles S.W. Bombay 17°

54'N 72 °27’E, (USNM 716676); Madras (BMNH 1953.1.30.190-—

4); Madras (BMNH 89.2.17.23-6); Tondi, Madras (NMW acc

55.158); Manapad (BMNH 1953.1.30.26); Mandapam, Gulf of

Mannar, 3m (ANSP 302433); Tuticorin (BMNH 1953.1.30.188—

89); S. of Krusadai I., Gulf of Mannar, 18m (ANSP 301196);

Andaman Islands (AM C.040397, C.040597); Andaman Islands

(NMW 55.158). SRI LANKA: Colombo (AM); Trincomallee, 29m

(AM); E., S., and N. coasts (AM C.309467); BMNH 1963491; Palk

Strait, 22m (AM); 9°34.5'N 80°39'E (AM C.309461); northern tip of

Eluvativu I., (ANSP 211085). BURMA: 57 miles N.W. Tavoy L.,

Anadamans Sea, 39m (ANSP 293254). THAILAND: Koh

Kahdat(AM C.031287); Phuket (USNM 661156); south end Light-

house I., Phuket, 5m (ANSP 286500); 25 miles N.N.W. Phuket, 42m

(ANSP 291734). MALAYSIA: Mersing Island, E. Malaya, 3.5m

(AM). SINGAPORE: BMNH acc. 2172; Singapore (AM C.3437);

NMW acc 55.158. INDONESIA: Kalimantan (AM C.309470).

PHILIPPINES: Corregidor, Bay of Manila, intertidal (BMNH;

Manila Bay, 18m (ANSP 281443); Manila Bay, 18m (ANSP 328655);

off Antonia I., E. Panay, 40m (USNM 293697); N. of Marinduque,

353m (USNM 295197); Lataau I., 33m (USNM 236058); Malampaya

Sound, Palawan, 25—35m (USNM 297395); off Tinkata I., Tawi

Tawi Islands, 30m (USNM 257677); VIETNAM: Vung Tau, Chilins

(ANSP 330797). AUSTRALIA: Western Australia, Shark Bay

(AM C.69245); Carnarvon, 7.6m (AM); Carnarvon (AM C.69270);

210 miles N. of Broome, 230m (AM); 110 miles N. of Melville

Island, Arafura Sea (AM); 72 miles N.W. of Dampier Island,110m

(AM 2405).

Cardiolucina siquijorensis Taylor and Glover, new species

Fig. 25

TYPE MATERIAL. Holotype, articulated specimen (Fig. 25b) USNM

812176. H=3.4 mm, L=3.5 mm. Paratypes, USNM 812177, 812169.

TYPELOCALITY. Siquijor, Solong-on, Siquijor I., Visayan Islands,

Philippines (09° 13'10"N 123° 27'30"E) depth 2-3 m.

SHELL DESCRIPTION. Shell very small, toa maximum height of 3.5

mm, outline circular H/L 0. 97, inflated, T/L 0.39, tumidity of single

valve to 1.3 mm. Shallow posterior sulcus with concentric lamellae.

Anterior sulcus absent. Lunule shallow, heart-shaped. Anterodorsal

area indistinct, posterodorsal area with concentric lamellae. Exte-

rior sculpture, fenestrate, of 1 1-12 broad, straight-sided, radial ribs.

Interspaces wide, but narrower than the ribs. Ribs are slightly

broader towards the anterior. About 14—16 thin, concentric lamellae,

project slightly above radial ribs. Hinge plate robust. Left valve with

single posterior lateral tooth, the dorsal edge of the socket extended

into a sharp rim (Fig. 22 e); two cardinal teeth, triangular in shape

and more or less equisized; single anterior lateral tooth. Right valve

with single anterior lateral tooth, a single central cardinal tooth and

a robust posterior lateral tooth. Ligament extends from beak to the

dorsal end of the posterior lateral tooth (Fig. 22 e). Inner margin with

12 coarse crenulations, and fine beading extending to the hinge

margins. Anterior adductor muscle scar elongate, not buttressed,

with only ventral tip detached from pallial line. Posterior adductor

muscle scar rounded. Pallial line continuous. Shell creamy white.

COMPARISON WITH OTHER SPECIES. This species is similar in size

and ornament to C. pisiformis but is distinguished by fewer and

broader radial ribs, the shallower lunule, the two equal-sized cardi-

nal teeth in the left valve and the single cardinal in the right valve.

115

HABITAT AND DISTRIBUTION. As for type locality.

MATERIAL EXAMINED. PHILIPPINES: Siquijor I., collected with

type material (USNM 812177, 812176, 812169).

Cardiolucina sp.

Fig. 26

In addition to the species described, we have four lots from the

Philippines consisting of rather badly preserved, disarticulated shells

and two rather better preserved juvenile shells. These are distinct in

a number of shell characters from the other Cardiolucina species.

Because the adult specimens are badly preserved and because

juveniles often differ considerably from adults, we are reluctant to

describe a new species based on this material.

The adult shells have maximum height of 10.8mm, with tumidity

of a single valve to 2.7mm and H/L 0.98, T/L 0.5. The posterior

sulcus is broad and shallow, the anterior sulcus absent. About 17-18

prominently scalloped, concentric lamellae, with more than 20 fine,

rounded, radial ribs, which do not cross the concentric lamellae. The

interspaces between the ribs are narrow, but relatively deep and the

intersection of the concentric and radial ornament produces a deeply-

pitted appearance. The lunule is heart shaped and relatively deep. In

the left valve, there is a single posterior lateral, two triangular

cardinal teeth, and a single posterior lateral. The inner margin of the

shell has about 25 crenulations.

This species is similar to C. semperiana but can be distinguished

by the lack of the anterior sulcus, the shallower posterior sulcus, the

pitted appearance of the shell surface and the more rounded

anterodorsal area.

MATERIAL EXAMINED. PHILIPPINES: off Sibugay I., E. of

Masbate, 198 m (USNM 292666); Daram Channnel, W. Samar, 48 m

(USNM 292830); off Matocot Point, W. Luzon, 400 m (USNM

295706); off Corregidor Light, 73 m (USNM 294200).

ATLANTIC OCEAN SPECIES

Cardiolucina lamothei (Dautzenberg, 1913)

Figs 27 a & b, 28

Lucina lamothei Dautzenberg, 1913: 100-101, pl. 3 figs 50-54.

Phacoides (Parvilucina) congoensis Thiele & Jaeckel, 1931: 220—

221, pl. 4 fig. 91.

Phacoides (Bellucina) lamothei (Dautzenberg) — Lamy 1920 : 215.

TYPE MATERIAL. Figured specimens of L. lamothei—two syntypes

from Tamara, Archipel de Los, Guinée, marked TYPE in

Dautzenberg’s handwriting (MNHN). Numerous other specimens

from localities mentioned by Dautzenberg (1913) are housed in the

Royal Belgian Institute of Natural Sciences (IRSNB inventory I.G.

10591). Our figured specimen is from a sample sent by Dautzenberg

to Tomlin (NMW 1955.158).

TYPE LOCALITY. West Africa, Tamara, Archipel de Los, Guinée.

SHELL DESCRIPTION. Shell solid, very small, height to maximum

of 4.0 mm. Subquadrate, inequilateral, slightly extended anteriorly.

Not inflated. Anterior sulcus shallow leading to marginal sinus.

Umbones are conspicuous, prosogyrate. Lunule very small, under-

neath beaks. Anterodorsal area indistinct and slightly convex.

Escutcheon with concentric lamellae. Exterior sculpture of 15-18

prominent, narrow concentric lamellae, which are slightly reflexed.

About 40 fine, radial ribs in the interspaces between concentric

116

J.D. TAYLOR AND E.A. GLOVER

Fig. 26

bars a = 400um, b = 500um, c=1mm.

lamellae. Hinge plate thick. Left valve with single, hooked anterior

lateral tooth, two cardinals and single posterior lateral tooth. Right

valve with similar. Indentations on both anterior and posterior

margins which correspond to the position of sulci. Inner margin with

about 40 crenulations. Anterior adductor muscle scar elongate, not

buttressed. Pallial line attached to lower third of anterior adductor

muscle scar. Posterior adductor scar reniform in outline. Pallial line

continuous. Shell colour grey-white.

COMPARISON WITH OTHER SPECIES. ‘This species is similar to C.

. rehderi but has a greater number of fine radial ribs.

HABITAT. Fine muddy sand from 4—200 m.

DISTRIBUTION. Senegal to Northern Angola (Fig. 28).

MATERIAL EXAMINED. AFRICA: Guinea, Los Islands (NMW

1955.158).

Cardiolucina rehderi (Britton, 1972)

Fig.27c &d

Parvilucina (Bellucina) rehderi Britton, 1972: 7-9, fig. 3.

Lucina (Parvilucina) rehderi (Britton) — Rios, 1994: 252.

TYPE MATERIAL. Holotype, USNM 208255, L = 4.2 mm, H = 4.2

mm, T = 4.1 mm (paired valves), also 9 paratypes.

a—-b, Cardiolucina sp. Corregidor, Philippines (USNM 294200); c, Cardiolucina sp Sibugay Island, Masbate, Philippines (USNM 29266). Scale

TYPE LOCALITY. Brazil, no further details.

SHELL DESCRIPTION. Shell small, height to 4.2 mm. Subcircular;

moderately inflated, maximum tumidity on a single valve to 1.9 mm.

Inequilateral, slightly extended anteriorly. Broad posterior sulcus.

Posterior dorsal area slightly convex, with distinct elevation of

concentric lamellae. Shallow anterior sulcus, anterior dorsal area

indistinct. Lunule shallow, heart-shaped. Exterior sculpture of about

30 indistinct radial ribs most clearly visible in the interspaces

between concentric lamellae, ribs separated by shallow, narrow

interspaces. Concentric lamellae,14—16, prominent, thickened la-

mellae often appear scalloped at position of intersection with radial

ribs. Hinge plate thickened. Left valve with single anterior lateral

tooth; two cardinal teeth and single posterior lateral. Right valve

with similar. Ligament short, extending about half distance from

beak to posterior lateral tooth. Inner margin with about 20 fine

crenulations. Anterior adductor muscle scar longer than wide but

relatively short with pallial line joining near ventral tip. Posterior

adductor muscle scar rounded. Shell colour white. A more extensive

description of this species is given by Britton (1972).

COMPARISON WITH OTHER SPECIES.

to C. lamothei.

This species is closely similar

HABITAT. Unknown.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

7 eRe **

" . ee &. % he ae

117

Fig. 27 Cardiolucina lamothei (Dautzenberg, 1913) and C. rehderi (Britton, 1972). a, C. lamothei Iles de Los, nr Conakry, Guinea (NMW 1955.158) Left

valve. P35.082; b, left valve interior; c, C. rehderi Britton, left valve, Paratype NMNH,; d, left valve interior. Scale bar a—d = 1.0mm.

Fig. 28 Geographical distribution of Cardiolucina lamothei

(Dautzenberg). Data from Dautzenberg (1913).

DISTRIBUTION. Known only from type material reported on by

Britton (1972).

FOSSIL SPECIES OF CARDIOLUCINA

Apart from the description of the type species C. agassizi, we have

not attempted a comprehensive review of the fossil record. How-

ever, the species listed below have the characters of Cardiolucina.

Other fossil species that we do not consider to belong the genus are

listed separately.

Phacoides (Parvilucina) ligatus Cossmann and Pissarro,

1904. (pages 17-18, pl. 7, figs 1-4).

Described from the Eocene (Lutetian) of Hauteville, Cotentin Pe-

ninsula, France. This small species has a thick shell and prominent

concentric lamellae with fine radial ribbing similar to that of

Cardiolucina agassizi. Chavan (1937: 207) first suggested that this

species should be classified as a Bellucina. On the basis of the

published figures, we consider that this is probably the earliest

known Cardiolucina.

Lucina nuciformis Tate, 1887b: 144 (figured in Tate, 1886,

p. 158, pl. 12 figs.10a & b).

118

This Miocene species from Blanche Point, Aldinga Bay, South

Australia, is very similar to the Recent Cardiolucina crassilirata

(Tate).

Linga (Bellucina) gonzalesi Shuto,1971 (pp. 30-31; Plate

1 figs 8,11,13—17; text fig. 6)

Described from the Lower Pliocene Cabatuan Formation of Panay

Island, Philippines. This species is well described and illustrated. It

has about eight prominent, radial ribs with a fenestrate ornament like

the Recent Cardiolucina eucosmia, C. pisiformis andC. siquijorensis.

The intersections of the radial ribs and concentric lamellae are

‘strongly granulated’, a feature found in none of the three Recent

species.

Lucina polli Icke & Martin, 1907. (p. 250, plate 18 figs 40,

40a)

Described from the Pliocene of Nias Island, Indonesia. From the

illustrations, this is a definite Cardiolucina species with fine radial

ribbing and prominent concentric lamellae, similar in form to C.

rugosa.

Cardiolucina agassizi var. regularior Sacco, 1904 (p. 90,

pl. 20 figs. 40-42) and var. constricta Sacco,1904 (p. 90 pl.

20 fig. 43).

Without a morphological analysis of C. agassizi populations it is

difficult to evaluate these names, but they would seem to be forms of

a variable species.

SPECIES EXCLUDED FROM CARDIOLUCINA.

Listed under original generic assignments as published.

Phacoides (Bellucina) actinus Dall, 1903: 1385, pl. 52 fig.

3. Also Woodring (1925: 126-127; pl. 17 figs 5-8).

Described from the middle Miocene Bowden Formation of Jamaica,

this species has 16—22 radial ribs and fine concentric lamellae.

However, the lunule is lanceolate and the hinge different from

Cardiolucina species. We consider that it may belong to the same

clade as Radiolucina amianta (Dall) (see below).

Phacoides (Bellucina) amiantus Dall, 1901: 826-7, pl. 39

fig.10.

Although Dall included this Caribbean species in Bellucina and

Bretsky (1976) used it as the basis of her diagnosis of the genus, it

differs in many features from the type species of Cardiolucina and

Bellucina. Britton (1972) discussed this at some length and pro-

posed the new generic name Radiolucina with P. amiantus as the

type species.

Phacoides (Bellucina) tuomeyi Dall, 1903: 1385, pl. 52

fig. 1.

Described from the late Miocene Duplin Marl of North Carolina,

this species is similar to the Recent Radiolucina amianta (Dall).

Phacoides (Bellucina) waccamawensis Dall,1903: 1385,

pl. 52 fig. 2.

Described from the Pliocene Waccamaw Formation of South Caro-

lina, this species is closely similar to the Recent Radiolucina ami-

J.D. TAYLOR AND E.A. GLOVER

anta (Dall) and was tentatively synonymised by Bretsky (1976).

Phacoides peritaphros Barnard, 1964: 476-477 fig. 22.

Syntypes: BMNH 1964264

Although Barnard regarded this southern African species as allied to

Cardiolucina lamothei (Dautzenberg), it lacks radial riblets, has a

shallow, open lunule and is more anteriorly extended. This species is

similar in many features to Cavilinga trisulcata (Conrad, 1841) (see

Olsson and Harbison 1953, pl.7 fig. 4), the type species of the genus

Cavilinga Chavan, 1937.

Lucina cancellaris Philippi, 1846: 21.

This is a West American twin species to the Caribbean Radiolucina

amianta. Keen (1971, p. 121) points out that ‘Dall unfortunately

assigned this species and its Caribbean twin L. amianta (Dall, 1901),

to the subgenus Bellucina, the type of which is an Indian Ocean form

only superficially similar, and American authors have been slow to

recognise the error.’

Lucina pulchella Lynge, 1909: 173, pl 3 figs. 13-15.

Lynge compared this species with L. seminula Gould, L. pisum

Reeve and L. semperiana Issel. We examined the type material

(Zoological Museum, Copenhagen) which has a different ribbing

pattern from any Cardiolucina and lacks posterior and anterior sulci.

This species may be an Epicodakia.

Lucina valida Smith, 1904: p.40 pl.3 fig.19.

This southern African species is very similar to and a probable senior

synonym of Phacoides peritaphros Barnard. Syntypes: BMNH

1903.12.19.1270-4.

Cardiolucina striatula var. taurotrigona Sacco, 1901: 90,

pl. 20 fig. 44-47.

Cardiolucina striatula var. ovatuloides Sacco, 1901 90 pl.

20 fig. 48.

Cardiolucina taurocrenulata Sacco, 1901: 90 pl. 20 fig. 50.

C. taurocrenulata var. aviculina Sacco, 1901: 90 pl. 20

figs 51 & 52.

Cardiolucina oligobliqua Sacco, 1901: 91 pl. 20 fig. 53.

All these taxa were described from the Miocene of Italy. They have

concentric ornament only and differ in shape from the type species.

In our opinion these taxa should all be excluded from Cardiolucina.

RELATIONSHIPS AMONGST THE SPECIES

In an attempt to establish relationships amongst the Cardiolucina

species a preliminary phylogenetic analysis using parsimony was

performed using a set of 12 shell characters. We used PAUP version

3.0, with further interpretation of characters with McClade version

3.04. Although it would have been desirable to include anatomical

characters these were available for only two species. We selected as

an outgroup the northeastern Pacific Parvilucina tenuisculpta (Car-

penter, 1864), the type species of the genus Parvilucina. Since the

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

o -~ &

a jana tae anh

[sth eof rian meh oe aa

TS Oe AGO! Ee IO er hS

a ee ek oe

Ss ts gut amy ie Cr cement item gee las

pisiformis

119

outgroup

3 = ~ =

2 a = = &

2 Ks) * g 5

ae) 0 be = A)

5 3 3 g 2 3

S = = 2 g oO

7) As} iy 1s) 1s) ~

6 3

Fig. 29 Strict consensus tree of Cardiolucina species derived from two equally parsimonious trees. Outgroup is Parvilucina tenuisculpta. Character state

changes at numbered internal branches. Branch 1: char. 1, state 2>1; char. 3 state 3>0; char. 6 state 2>1. Branch 2: char. 3 state 0>2; char. 8 state 0>1;

char. 11 state O>1. Branch 3: char. 11 state 1>2. Branch 4: char. 7 state 1>0. Branch 5: char. 4 state 0>1; char.5S state O>1; char. 6 state 1>0; char. 12 state

2>0. Branch 6: char. 9 state 0>1. Branch 7: char. 1 state 1>0; char. 2 state 0>2. Branch 8: char. 12 state 2>1. Branch 9: char. 2 state 2>1. Branch 10:

char.10 state O>1.

Table 1. Characters and character state codings used in the phylogenetic analysis.

1. Posterior sulcus. This is an indentation in the posterior shell seen on

the outer surface extending from the umbo to the shell margin. It is present

in all Cardiolucina species, deeply incised in some (Fig. 20d), shallow in

others (Fig. 2a) and weak and poorly-defined in the outgroup.

States: 0 - deep prominent;1 - shallow; 2 - weak

2. Anterior sulcus. This is a similar indentation at the anterior end of the

shell. This is absent in some species, but where present (Fig. 20) never as

prominent as the posterior sulcus.

States: 0 - absent; | - narrow; 2 - broad

3. Lunule. Many species have a heart-shaped, shallow lunule (Fig. 20c) ,

but in C. eucosmia it is deeply excavated. In other species the lunule is

very small and tucked under beak (Fig.2) and in the outgroup it is

lanceolate.

States: 0 - shallow, heart-shaped; | - deeply excavated; 2 - tiny under

umbo; 3 - thin lanceolate

4. Radial ribs - number

States: 0 - less than 20; 1 - more than 20

5. Radial rib - shape

Some species have prominent ribs with straight sides and separated by

broad interspaces (Figs. 2,3). In others, the ribs are closer together with

low rounded profiles and narrow interspaces (Figs. 7,10).

States: 0 - rounded, narrow interspaces; | - straight-sided broad interspaces

6. Radial rib intersection with concentrics.

In some species, ribs appear to cross concentric lamellae without

interruption (Fig. 2d) whilst in others the ribs terminate at the intersection

with major concentric lamellae (Fig. 7d). In the outgroup the radial ribs

are extremely weak.

States: 0 - strong, cross concentrics ; | - weak, terminate at concentrics; 2 -

very weak

7. Concentric lamellae - number

Concentric lamellae are a prominent feature of many Cardiolucina species.

States: 0 - less than 20; 1 - more than 20

8. Shape of anterior cardinal tooth in left valve

The majority of species have a triangular wedge shaped tooth (Fig. 18b)

but in some species the anterior cardinal is a hook-shaped projection (Fig.

3109):

States: 0 - wedge; 1 - hooked

9. Posterior cardinal of left valve

Most species have a relatively thin tooth (Fig. 7c ), but in C. pisiformis ,

for example, it is much thicker (Fig. 14c ).

States: 0 - thin; 1 - thick

10. Denticles on cardinal teeth

This refers to small denticles seen at high magnification along the edges of

the posterior cardinal (Fig. 12d).

States: 0- absent; 1 - present

11. Hinge buttress

The hinge may be projected ventrally below the cardinal teeth to form a

rounded buttress (Fig 3c).

States: 0 - absent; | - slight; 2 - prominent

12. Margin crenulations - number

These are seen on the internal ventral margin of shell. These were counted

around the shell margin, fine beading not included.

States: 0 - coarse (n = 10-20); 1 - medium (n = 21-30); 2 - fine (31 + ).

120

Table 2 Matrix of Cardiolucina species and the outgroup Parvilucina

tenuisculpta and their character states. Character coding as in Table 1.

OQ 7 9 10 Ii

—

agassizi 0

australopilula l

civica 1

crassilirata 1

eucosmia ]

pisiformis 1

quadrata 1

rugosa 0

semperiana 0

siquijorensis 1

lamothei 0

rehderi 0

P.tenuisculpta 2

SnnowrrHrooracoco

WBOSCSCCONOHNNNO |] w

Be ROR RF RF OOH OF

SeoOOHMOoOCOoOKRHOOHO|U

NeEH Oe Ee HooHrHor

SOOCOCOOFOOHKHH OO]

SeooOrooorcecoce

SOSONuRH HK OOHKoCoOCSO

eecooocoonooeene

NHENOHKPHK RB OCONNHE=

Soo oom oot 1S StS

original designation by Dall (1901), Cardiolucina (as Bellucina) has

been widely listed as a subgenus of Parvilucina and the shells have

some features in common (descriptions in Hickman 1994). More-

over, there are some similar anatomical features, P tenuisculpta

having a visceral lobe resembling that found in C. semperiana and

C. australopilula, as well as the connection of the posterior ctenidia

to the mantle and the lack of a fused inhalant aperture (Reid and

Brand, 1986). The characters and their states are listed in Table 1.

The distribution of the various character states amongst the

Cardiolucina species and the outgroup is shown in Table 2.

Phylogenetic analysis using this matrix produced 2 equally parsimo-

nious trees of 26 steps with a consistency index of 0.731 and a

homoplasy index of 0.269. A strict consensus tree is shown in Fig.

29. The differences in the 2 trees resulted from the grouping of

C. civica and C. crassilirata as a single clade or as a trichotomy with

the C. australopilula /C. quadrata clade.

The analysis suggests that there are three distinct clades of

Cardiolucina species (Fig. 29). Clade 1 contains C. australopilula,

Table 3 Summary of shell characters in Cardiolucina species

J.D. TAYLOR AND E.A. GLOVER

crassilirata, civica and quadrata. Apomorphies which define this

clade at Branch 2 are; the tiny lunule, the hooked-shaped, anterior

cardinal tooth and the hinge buttress. Clade 2, separated at Branch 4

by a single apomorphy of having less than 20 concentric lamellae,

contains two major branches; that comprising the type species C.

agassizi and the species around C. semperiana. (Branch 7) and

another branch (Branch 5) containing C. eucosmia, pisiformis and

siquijorensis. This latter clade is defined by four apomorphies;

having less than 20 radial ribs, which are straight-sided and cross the

concentric lamellae and also possess coarse marginal crenulations.

The C. agassizi — lamothei clade is defined at Branch 7 by the

prominent posterior sulcus and a broad anterior sulcus.

This hypothesis of relationships should be regarded as tentative

because it is based on shell characters alone. Nevertheless, it is open

to falsification by additional or alternative character sets.

GEOGRAPHICAL DISTRIBUTION

Two species, Cardiolucina lamothei and C. rehderi are found on

opposite sides of the Atlantic, whilst the other nine are distributed

largely within the tropical Indo-West Pacific Province. One of these,

C. semperiana, has a wide range from the Gulf of Suez to Philip-

pines whilst the rest have more restricted distributions within the

central Indo-Pacific. Four species are endemic to Australia; C.

eucosmia, C. pisiformis and C. australopilula to the western and

northern coasts and C. crassilirata to the temperate southern coast.

Cardiolucina civica occurs from Japan through Indonesia to the

Andaman Islands in the west and Fiji in the east. C. quadrata is

restricted to deeper water around the Philippines and Indonesia.

Apart from C. rehderi, which is known from only one vague locality,

the species with the most limited range is C. siquijorenis from

Siquijor Island in the Philippines. All the species are largely tropical

in distribution, except for C. crassilirata from southern Australia.

Lunule

Sulcus Marginal small, under _heart-

Species H L T Radial ribs Concentric lamellae post. ant. crenulae beak shaped

agassizi 74 71 2.7 ¢.25 faint, closely c.20 width of interspaces deep narrow 28 shallow

spaced extremely variable

australopilula 13.8 12.7 7.5 c.20 interspacesalmost c.34-36 low, narrow shallow none 21 a

as wide as ribs

civica 10.6 104 45 >30 fine, closely >25 conspicuous shallow none >60 =

spaced

crassilirata Hi) tt3} >40 fine, closely >30 fine, clustered shallow none 31 as

spaced in groups of 4-6

eucosmia 7.8 7.7 3.8 c.12 broad, with c.13-14 low v.deep narrow 12 v. deep

wide interspaces

pisiformis 44 44 2 13-17 low, widely c.20 narrow shallow none 13 deep

spaced

quadrata 12.9 13.6 5.3 ¢.25 fine,closely 25-30 low, broad shallow none 25 *

spaced

rugosa 331 3.2 1.2 >25 faint, closely c.20 prominent deep narrow 20 shallow

spaced

semperiana Well 7.5 2.7 20-30 low, closely 15-24 interspaces variable deep narrow, 20-21 shallow

spaced in width ribbed

siquijorensis 3.5 3).5) 13 12 broad 14-16 thin shallow none 12 shallow

lamothei 4 310) ae >40 fine, closely 15-18 prominent deep broad 40 s

spaced

rehderi 42 42 1.9 >30 fine, faint 14-16 prominent deep broad 20 3

closely spaced

H=height, max mm, L=length, max mm, T=tumidity, max mm.

ANATOMY AND SYSTEMATICS OF CARDIOLUCINA

The phylogenetic analysis shows that Cardiolucina lamothei and

C. rehderi are closer to the C. semperiana clade of species than to the

C. australopilula or C. eucosmia clades. This suggests that the clade

containing C. lamothei-rehderi and C. semperiana had a widespread

Tethyean distribution prior to the (late Miocene) closure of the

connection between the Indo-Pacific and Atlantic Oceans. This is

confirmed by the occurrence of C. agassizi in the Middle Miocene of

south west Europe. The C. australopilula clade is restricted in

distribution to the central Indo-Pacific. The Miocene species C.

nuciformis Tate from South Australia is similar to the Recent C.

crassilirata from the same area suggesting a long occupancy.

CONCLUSIONS

In this revision of the systematics of Cardiolucinawe have redefined

the genus and its type species and recognised eleven living species

from the tropical Atlantic and Indo-W. Pacific provinces. The previ-

ous lumping of many taxa into C. semperiana underestimated

species diversity. Table 3 provides a summary of the characters of the

recognised species as an aid to identification. Anatomical characters

are known for only two species, C. australopilula andC. semperiana.

Comparison with other genera is difficult because of the lack of

information, but Parvilucina has some similar characters, such as

the lack of mantle fusion at the posterior inhalant area, the connec-

tion of the ctenidia to the posterior mantle and the large visceral lobe

(Reid and Brand, 1986). Further work is needed to establish the

distribution of these characters in other lucinid taxa.

Although the earliest Cardiolucina would seem to be the Eocene

species C. ligata (Cossmann & Pissarro), the fossil record, apart

from C. agassizi which is common in the middle Miocene of Europe,

is rather scanty. The only species we can confirm are Cardiolucina

nuciformis (Tate) from the Miocene and, from the Pliocene, C.

gonzalesi (Shuto) and C. polli (Icke & Martin). This paucity of

records is unsurprising because, even in Recent faunas, these small

lucinids are neglected.

The problems encountered during this systematic revision reflect

the current state of systematics within the family Lucinidae. Many

genera are defined only by a small number of shell characters, type

species of genera are often inadequately described and, moreover,

their characters are often ignored by systematists. Species diversity

is often greater than realised because taxa have been shoe-horned

into a small number of ‘well-known’ names. Relationships between

genera are generally unknown and rigorous phylogenetic analyses

are lacking. Bretsky (1970, 1976) produced a phenetic/phylogenetic

analysis of lucinid genera, with possible phylogenies for her various

lineages. Unfortunately, there are problems with these phylogenies

because she lumped together distinct genera and confused type

species, for example, using for Bellucina characters of Radiolucina

amianta instead of the type species B. eucosmia. Moreover, there is

no anatomical or biological information available for the vast major-

ity of species, especially those from the Indo-Pacific province.

Recent studies on the chemosymbiosis of lucinids have focused on

a small number of species from the Caribbean and temperate north

Atlantic and Pacific (reviewed in Fisher, 1990, Le Pennecet al.,1995),

but biological information from these studies has yet to be integrated

into lucinid systematics. Systematic and phylogenetic studies should

be providing the framework to test hypotheses concerning the

evolutionary history and radiation of the Lucinidae, but, at present,

these are inadequate.

121

ACKNOWLEDGEMENTS. Ian Loch, Jerry Harasewych, Tom Schiotte,

Graham Oliver & Alison Trew, Philippe Bouchet & Rudo von Cosel, Dick

Kilburn, Robert Moolenbeek, Fred Wells & Shirley Slack-Smith, Hiroshi

Saito and John Slapcinsky generously provided access to material and loan of

specimens in their care. Thierry Backeljau kindly sent information about the

Dautzenberg collection. Philippe Bouchet & Rudo von Cosel provided

access to unpublished information and commented on the manuscript. We

thank Graham Oliver for discussion and nomenclatural advice. Additionally

we are grateful to David Cooper for the thin sections of C. australopilula,

Harry Taylor for fine macrophotography, Nick Hayes for printing of SEM

photographs and Alex Ball for translating some descriptions.

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Wissenschafftliche Ergebnisse der Deutschen Tiefsee-Expedition, 21: 1-268.

Woodring, W.P. 1925. Contributions to the geology and paleontology of the West

Indies. Miocene mollusks from Bowden, Jamaica. Pelecypods and scaphopods.

Carnegie Institute of Washington Publication, 366: 1-222.

Yokoyama, M. 1927. Fossil Mollusca from Kaga. Journal of the Faculty of Science

Imperial University of Tokyo. Section II (Geology), 2 (part 4): 165-182.

Bull. nat. Hist. Mus. Lond. (Zool.) 63(2): 123-128

Issued 28 November 1997

A new species of water mouse, of the genus

Chibchanomys (Rodentia, Muridae,

Sigmodontinae) from Ecuador

PAULINA D. JENKINS ~

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, U.K.

ADRIAN A. BARNETT

School of Life Sciences, Roehampton Institute, West Hill, London SW15 5SN, U.K.

SyNopsIs. A new species of the rodent genus Chibchanomys is described from Ecuador on the basis of external and cranial

morphology. A phylogenetic analysis is used to assess the relationship of the new species to other ichthyomyine taxa.

INTRODUCTION

The New World rodents currently placed in the large and complex

subfamily Sigmodontinae (sensu Carleton & Musser, 1984) include

amorphologically and ecologically distinctive group of semi-aquatic

South American genera (Voss, 1988), for which it is convenient here

to use the name ichthyomyines. In a monograph of ichthyomyine

rodents, Voss (1988) included five genera: Ichthyomys Thomas,

1893 (four species), Anotomys Thomas, 1906a (monotypic), Rheomys

Thomas, 1906b (four species), Neusticomys Anthony, 1921 (four

species) and a new genus, Chibchanomys Voss, 1988 for an enig-

matic species of uncertain generic affinity. Chibchanomys trichotis

(Thomas, 1897) was originally placed in Jchthyomys but was subse-

quently assigned to Rheomys (see Tate, 1932; Cabrera, 1961), then

to Anotomys (see Handley, 1976). Subsequent to the revision by

Voss, an additional ichthyomyine species, Neusticomys mussoi Ochoa

& Soriano, 1991, has been described.

During the course of several zoological surveys of Las Cajas

Plateau, Ecuador from 1981 to 1987, five specimens of an undescribed

species of ichthyomyine rodent were captured. Observations were

made on two of these animals, which subsequently escaped, while

three specimens were donated to The Natural History Museum.

Another specimen was filmed for the 1992 BBC National Geo-

graphic wildlife film ‘Avenue of the Volcanoes’ (Jim and Theresa

Clare, personal communication). On the basis of external and

craniodental characters the study specimens agree most closely with

the generic diagnosis of Chibchanomys but are also sufficiently

similar in some features to Neusticomys to warrant a phylogenetic

analysis.

MATERIALS AND METHODS

The description of the new species is based on conventional morpho-

logical characteristics and the terminology used follows Voss (1988).

Specimens were measured using dial calipers, with all measure-

ments provided in millimetres. The skeletal elements remaining in

the skins were observed by means of X-rays.

A parsimony analysis (PAUP Version 3.0) was carried out to

determine the position of the new species relative to other

ichthyomyines. Details of the eighteen characters used, listed

below, are given more fully in Voss (1988 pages 440-442); the

hypothesised primitive state, using the criteria defined by Voss, is

scored as ‘0’.

. Pelage: glossy (0); dull (1).

. Ventral pelage countershaded: absent (0); present (1).

. Tail: unicolored (0); bicolored (1).

. Philtrum: present (0); absent (1).

. Pinnae: large, visible above fur (0); small, concealed in fur (1).

. Superciliary vibrissae: present (0); absent (1).

. Plantar pads of manus: hypothenar pad separate, not fused with

third interdigital pad (0); hypothenar and third interdigital pads

fused (1); hypothenar and thenar pads fused respectively with

adjacent third and first interdigital pads (2).

. Fringing hairs on pes: weakly developed (0); well developed (1).

9. Lower third molar: entoconid-hypoconid cusp pair distinct (0):

m3 peglike, entoconid-hypoconid cusp pair absent or reduced to

a small conule (1).

10. Nasal bones: long, produced anteriorly beyond premaxillae (0);

short, truncated behind premaxillae (1).

11. Supraorbital foramina: on the lateral surface of the frontals,

within orbital fossae (0); on the dorsal surface of the frontals

between the orbital fossae (1).

12. Carotid arterial supply (see Voss, 1988 page 296): pattern 1 (0);

pattern 2 (1); pattern 3 (2).

13. Orbicular apophysis of maleus: present (0); absent (1).

14. Metatarsal configuration: WI>IV >>> V>1(0);IV>I1> 1H,

V>I1(1).

15. Omohyoid muscle: present (0); absent (1).

16. Gastric glandular epithelium: present (0); restricted (1).

17. Gall bladder: present (0); absent (1).

18. Bacular cartilage: tridigitate, medial digit lacking a calcified

centre (0); single digit (1); tridigitate, medial digit grossly

swollen with calcified core (2).

ADM fLWND —

The character states for the new species were assessed by PJ but the

character state assessments for the other taxa were taken directly

from Voss (1988 Table 45, page 441). Characters of the visceral and

reproductive systems (characters 15—18 above) were unobservable

in the new taxon because of the lack of whole bodies. In an initial

analysis, all character states were unordered; in a second analysis,

multistate characters were ordered (as by Voss): 0 ~ 1 — 2 for

characters 7, 12 but also for character 18. For character 18, the order

recommended by Voss (1988) was 0 — 1; 0 — 2, a sequence not

readily handled by the analysis and affecting only one generic group

124

(Rheomys); Voss (personal communication) recommended an alter-

native ordering of this character (1 > 0 — 2) so in a third analysis,

characters 7 and 12 were ordered as above and character 18 by this

alternative.

ABBREVIATIONS USED IN THE TEXT

BMNH - The Natural History Museum [formerly British Museum

(Natural History)]

M1, M2, M3 respectively first, second and third upper molars

ml,m2,m3 ___ respectively first, second and third lower molars

ABBREVIATIONS USED FOR TAXA IN THE

PAUP ANALYSIS:

Ale Anotomys leander Thomas, 1906a

Ctr Chibchanomys trichotis (Thomas, 1897)

Cor Chibchanomys undescribed species

Thy Ichthyomys hydrobates (Winge, 1891)

Ipi Ichthyomys pittieri (Handley & Mondolfi, 1963)

Itw Ichthyomys tweedii Anthony, 1921

Nmo Neusticomys monticolus Anthony, 1921

Nve Neusticomys venezuelae (Anthony, 1929)

Rme Rheomys mexicanus Goodwin, 1959

Rha Rheomys raptor Goldman, 1912

Rtt Rheomys thomasi Dickey, 1928

Run Rheomys underwoodi Thomas, 1906b

RESULTS

Chibchanomys orcesi, sp. nov.

HOLOTYPE. BMNH 82.816, adult male, skin and skull; collectors’

number 148; collected 22 August 1981 by members of the Oxford

Expedition to Las Cajas from Lake Luspa, Las Cajas, Provincia

Azuay, Ecuador, 02°50'S 79°30'W, altitude 3700m.

PARATYPES. BMNH 82.815, adult male, skin and skull; collectors’

number 146, other details as for the holotype. BMNH 84.349, adult

male, skin and skull; collectors’ number 78; collected 7 August 1983

by members of the Combined Universities Expedition to Ecuador

1983, from Lake Llaviucu, Zorracucho Valley, Las Cajas, Provincia

Azuay, Ecuador, 02°51'S 79°O1'W, altitude 3100m.

DIAGNOSIS

An ichthyomyine species belonging to the genus Chibchanomys in

the following combination of features. Dorsal pelage dull; small

pinnae concealed in pelage of head; tail longer than head and body;

manus with five separate plantar pads; hindfoot broad with well

developed fringing hairs; supraorbital foramina open laterally within

orbits; carotid circulation pattern 1.

Differing from Chibchanomys trichotis in the following charac-

ters. Rhinarium light brown; philtrum present; nasals medium in

length, barely projecting anterior to premaxillae; orbicular apophy-

sis of maleus present; upper incisors slightly inclined medially; M3

and m3 reduced in size; anteromedian flexid absent or barely

indicated on anteroconid of m1; metatarsals I] >IV > IL>V>L.

DESCRIPTION

Tail subequal to or slightly longer than head and body (see Table 1

P.D. JENKINS AND A. BARNETT

Table 1 External and cranial measurements of Chibchanomys trichotis

and C. orcesi. Dimensions given as mean, plus or minus standard

deviation, followed by range, with sample size in parentheses.

C. trichotis _C. trichotis_ C. trichotis C. orcesi

Venezuela Colombia Peru Ecuador

Head and body 113.5 +5.50 105 + 1.63

length 105-120 (4) 125 (1) 102 (1) 103-107 (3)

Tail length 126.8 + 7.36 113.3 +6.18

115-133 (4) 131(1) 123 (1) 108-122 (3)

Hindfoot length 31.8 + 1.09 22 +2.16

30-33 (4) 30) 3312)) “3iha) 19-24 (3)

Ear length Uz WS) WLS) Sey

6-10 (4) 8 (1) 6 (1) 9.5-14 (3)

Weight (in grams) - - - S72 2-83

3541 (3)

Ratio of tail 1.12 + 0.02 1.08 + 0.06

length to head 1.10-1.15 (4) 1.05 (1) 1.21 (1) 1.02—1.16(3)

and body length

Ratio of tail 4.9+0.12

length to condylo- 4.7—5.0 (4) Sel) 5.2 (1) 4.3, 4.7 (2)

incisive length

Ratio of hindfoot 0.28 + 0.00 0.21 + 0.02

length to head 0.28-0.29 (4) 0.26 (1) 0.30 (1) 0.18—0.23 (3)

and body length

Condyloincisive 25.7 +1.01

length 24.3-26.9 (4) 25.9 (1) 23.8 (1) 23.0, 24.6 (2)

Diastema Length 6.3 + 0.54 - 33/5029

5.7-7.0 (3) 5.7 (1) 5.0-5.7 (3)

Length of upper 4.4+0.08 4.2 + 0.05

molars 43-45 (3) 4.4(1) 4.2 (1) 4.14.2 (3)

Incisive foramina 5.0 + 0.27 4.5 + 0.09

length 4.6-5.3(4) 5.1 (1) 4.5 (1) 4.44.6 (3)

Breadth of 1.2+0.11 1.4+ 0.05

incisor tips 1.0-1.3 (4) cl.2,1.3(2) 1.0(1) 1.4-1.5 (3)

Breadth of incisive 1.9+0.15 2.1 + 0.09

foramina 1.7-2.1 (4) 2.4 (1) 1.9 (1) 2.0-2.2 (3)

Breadth of 3.1+0.29 2.2 + 0.08

palatal bridge 2.7-3.4 (3) - PHO) ((l)) 2.12.3 (3)

Nasal length 9.0 + 0.41 9.3 + 0.29

8.5-9.5 (3) 8.1 (1) 8.5 (1) 8.9-9.6 (3)

Nasal breadth 3.0 + 0.05 3:2 O05

2.9-3.0(4) 3.0(1) 2.8 (1) 3.1-3.2 (3)

Interorbital breadth 4.7 + 0.13 4.6 + 0.08

45-4.8(4) 4.9(1) 4.3 (1) 4.54.7 (3)

Zygomatic breadth 13.4 + 0.66

12.3-14.1 (4) c13.9 (1) 11.7 (1) c12.8 (1)

Braincase breadth 13.4+0.27

13.0-13.7 (4) 13.8 (1) 12.4 (1) 1.9, 12.0 (2)

Ratio of inter- 0.35 + 0.02

orbital breadth to 0.33-0.37 (4) 0.36 (1) 0.35 (1) 0.38, 0.39 (2)

braincase breadth

Breadth of 1.1+0.05 1.1+0

zygomatic plate 1.0-1.1 (4) 1.0,1.2(2) 1.0(1) 1.1 (3)

Breadth of first 1.6+0.05 1.3+0.05

upper molar 1.5-1.6 (3) 1.5 (1) 1.4 (1) 1.3-1.4 (3)

Height of upper 4.6 + 0.33 4.3-4.5 (3)

incisor 4.1-5.0(4) 5.3 (1) 3.9 (1) 44+ 0.08

Depth of upper 1.3+0.12 1.2-1.3 (3)

incisor 11-144) 12) 1.0 (1) 1.3 +0.05

Breadth across 74+0.18

occipital condyles 7.2—7.6 (4) 7.6 (1) TeSialy) 6.6, 6.7 (2)

A NEW SPECIES OF CHIBCHANOMYS

Fig. 1 Live specimen of Chibchanomys orcesi.

Fig. 2 Skull of Chibchanomys orcesi (BMNH 1982.816) from left to right in dorsal, ventral and lateral view.

126

for measurements). Pelage soft, dense and woolly, dark brownish

grey dorsally, light grey ventrally; tail greyish brown, densely

haired, grey brown and brown hairs predominate proximally, with a

proportional increase of buff and cream hairs distally, extending

beyond tip in a short pencil. Distal portion of muzzle light grey in

young adults, cream in older individuals (age based on degree of

dental wear); rhinartum light brown in dry specimens; philtrum

present; conspicuous silvery-grey mystacial vibrissae present. Pin-

nae small, concealed by pelage; region of more-or-less conspicuous

light grey hairs ventro-lateral to pinnae. Manus with three interdigital

and two carpal pads. Well developed fringe of stiff hairs on margin

of metatarsus and digits of pes; claw of fifth digit extends beyond

first interphalangeal joint of fourth digit; claw of first digit reaches

midway along first phalange of second digit. See Fig. 1 for external

features visible in a photograph of a live specimen.

Skull (see Fig. 2) with moderately long nasals, overlapping nasal

orifice to conceal incisors in dorsal view but barely projecting

beyond premaxillae; rostrum short and narrow, naso-lacrymal cap-

sules evident in dorsal view; interorbital region moderately narrow

relative to braincase breadth (0.38, 0.39 (n = 2); frontals slightly

inflated, braincase moderately broad and long; posterior border of

incisive foramina between anterior roots of M1s, palatal foramina lie

between posterior roots of M1s; bullae slightly inflated; orbicular

apophysis of maleus present. Carotid circulation pattern 1, based on

osteological features (see Voss, 1988: 298).

Upper incisors moderately narrow, anterior enamel surface pale

buff, slightly inclined medially. No anteroloph on M1; small

posteroloph on M2; M3 small, protocone and paracone evident in

unworn dentition, posterior conule absent. Anteroconid of m1 sim-

ple or with slight indication of anteromedian flexid; no anterolophid

on m2; small posterolophids on ml and m2, small mesolophids

present or absent; m3 small, with small posterior basally positioned

conulid.

Metatarsal proportions: third metatarsal slightly longer than fourth,

fourth longer than second; all three far longer than first and fifth;

fifth longer than first. Configuration: II] 21V > Il > V >I.

ETYMOLOGY

This species is named in honour of Professor Gustavo Orcés, a

pioneer of Ecuadorian mammalogy. He was of great help to AB with

fieldwork organisation in Ecuador, and his kindness and knowledge

were a source of inspiration.

DISTRIBUTION AND ECOLOGY

Known only from Las Cajas Plateau, Ecuador, where specimens

have been recorded from three localities: Lake Luspa, Lake Llaviucu,

Zorracucho Valley and Lake Torreadora. All specimens were trapped

in close proximity to fast-flowing streams at altitudes ranging from

3100m to 4000m, in high-altitude moorland vegetation (paramo)

(see Barnett, 1992). For more precise details of the habitat at each

site and notes on diet see Barnett (1997).

COMPARISON WITH C. TRICHOTIS

The new species is similar in external appearance to C. trichotis,

except that the pelage is paler and slightly harsher, and the rhinarium

is light brown in dry specimens of C. orcesi, black in C. trichotis. A

philtrum is present in C. orcesi but absent in C. trichotis.

Chibchanomys orcesiis smaller in external size and averages smaller

in cranial size than all known specimens of C. trichotis, with the

exception of the single specimen from Peru (see below for com-

ments on the status of this specimen). Both species of Chibchanomys

are similar in external proportions, except that the hindfoot is

proportionately shorter in C. orcesi (see Table 1). The metatarsal

P.D. JENKINS AND A. BARNETT

configuration differs in the two species: IV > III > Il=V>Iin C.

trichotis; W121V > IL >> V > Lin C. orcesi. The two species differ in

the following cranial features: while the nasals of both species are of

comparable length, those of C. orcesi are slightly broader and barely

project anterior to the premaxillae, unlike those of C. trichotis,

which project anteriorly and conceal the incisors and nasal orifice in

dorsal view. In lateral view, the globose braincase of C. trichotis

rises abruptly in the frontal region, unlike the narrower and less

inflated braincase of C. orcesi; the braincase is slightly broader and

the breadth across occipital condyles is greater in C. trichotis (see

Table 1). The orbicular apophysis of the maleus is present in C.

orcesi but absent in C. trichotis. The upper incisors of C. orcesi are

less delicate and slightly broader than those of C. trichotis (see Table

1), the anterior enamel surface of C. trichotis is cream coloured and

not medially inclined unlike C. orcesi. The third upper molar is

smaller relative to M1 and M2, and m3 is smaller relative to m1 and

m2 in C. orcesi than in C. trichotis. The anteromedian flexid is

absent or barely indicated on the anteroconid of m1 in C. orcesi but

present in C. trichotis, dividing the anteroconid into small but

distinct lingual and labial conulids. The posterior conulid of m3 is

positioned more basally in C. orcesi than in C. trichotis (when

present).

According to Voss (1988) the young specimen that he identified as

C. trichotis from Peru differs from the northern specimens of C.

trichotis in several features: the braincase is much less inflated, the

occipital condyles are slightly broader, the bullae are somewhat

smaller and an indistinct philtrum is indicated; features that resem-

ble those of the new species. Voss mentioned that these differences

might indicate that southern populations of Chibchanomys are

phenotypically distinctive from their northern counterparts. Unfor-

tunately it has not proved possible to examine the Peruvian specimen,

although information on it was kindly provided by Mark Hafner

(personal communication). It is possible that the Ecuadorian and

Peruvian specimens are conspecific but additional material and

more extensive comparisons are required to elucidate the status of

the latter specimen.

COMPARISON WITH OTHER ICHTHYOMINE GENERA

Chibchanomys is readily distinguished from Anotomys, Ichthyomys

and Rheomys (see Voss, 1988). Chibchanomys and Neusticomys

differ from other ichthyomyines in showing carotid arterial circula-

tion pattern | and in the distribution of the glandular epithelium

around the stomach. Chibchanomys differs from Neusticomys in

having small pinnae concealed in the pelage (pinnae obvious in

Neusticomys); ventral countershading present (absent in

Neusticomys); tail longer than head and body (tail shorter than head

and body in Neusticomys); the hindfoot is broader with longer digits

and the fringing hairs are well developed (narrower with shorter

digits and less developed fringing hairs in Neusticomys). The new

species does however share several features with Neusticomys which

are not exhibited by C. trichotis, such as the similar metatarsal

configuration and presence of a philtrum, while the orbicular apo-

physis of the malleus is also present in some species of Neusticomys.

RESULTS OF THE PHYLOGENETIC ANALYSIS

There is evidence in support of the ichthyomyines as a monophyletic

group of the subfamily Sigmodontinae (sensu Carleton & Musser,

1984) (see Voss, 1988). In contrast, evidence in support of the

monophyly of the Sigmodontinae is lacking and a tribal level

classification of this subfamily, while convenient in many respects,

is unsatisfactory from a phylogenetic point of view, making difficult

the choice of satisfactory outgroups for phylogenetic analyses (see

Voss, 1988: 436-438, 1991: 33-37; Carleton & Musser, 1989: 53—

55; Voss & Carleton, 1993: 21-22). The necessity of making such a

A NEW SPECIES OF CHIBCHANOMYS

Table 2. Matrix showing character state distributions among 12

ichthyomyine species (for details see text and Voss, 1988). The character

state assessments for all taxa other than C. orcesi were taken directly

from Voss (1988 Table 45, page 441). Character states for the new

species were assessed by PJ except for those of the visceral and

reproductive systems (characters 15—18) which were unobservable in

the new taxon, so scored as ‘?’.

Characters

eee a ee Ome, (o 00) eNO) eats 4 tL 6, 17) 8

Taxa

Ale

Ctr

Cor

Thy

Ipi

Itw

Nmo

Nve

Rme

Rha

Rtt

Run

MSM S) (SS) (SSS Siete

St oo ee ee

Se) SSS) (Sa) La aay (=)

eoOoorc”joq°”ccocdmh6cOlULcOlmrR hE

eS Sa CS) ne

a a)

NrreNocCcC oOo So co -

oon ee ee

SiO SS 1S (Si oS ic

Oe OS OO On = OOS

NNNNOCONNN COCK

(SS = SS SV) (Se) fe)

—— eS eS OOKe Se HS Oe eS

SESS) SW SiS ays) (SS) a

— RO OOO RR SE IO

Oo Oo © © 6 Oi = 3S

NNONNOCCOCOCyN O-

choice has been avoided in the current study, since it is aimed at

determining the level of affinity of the new taxon to other

ichthyomyines, rather than seeking to add any new dimension to the

phylogenetic status of the ichthyomyines as a group. Instead an

hypothetical outgroup was constructed in which all character states

were assessed as primitive, which was used to root the trees.

Using branch and bound algorithms, a search was made of the

character data summarised in Table 2. In the analysis in which all

character states were unordered, the length of the shortest tree was

equal to 32 character state transformations and six trees were

retained. In each of the other analyses (with the multistate characters

7 and 12 ordered, and character 18 varyingly ordered) the length of

the shortest tree was equal to 33 character state transformations but

127

only three trees were retained. The variation in treatment of charac-

ter 18, was not considered to be particularly important in this study,

since character state 2 is exhibited only by taxa of the genus

Rheomys. In both of the latter analyses C. orcesi and C. trichotis are

non-monophyletic in all three trees and also in the semistrict consen-

sus of these trees (see Fig. 3). The only evidence of a monophyletic

generic grouping shown in the semistrict consensus tree is for

Ichthyomys, and this tree is similar in most respects to the most

parsimonius hypothesis of ichthyomyine relationships shown by

Voss (1988: Fig.88).

DISCUSSION

There is obvious conflict in that the results of the phylogenetic

analysis do not support the generic classification currently in use.

The morphological data is sufficiently persuasive to conclude that,

on the available material, the new taxon is correctly attributed to the

genus Chibchanomys as currently construed.

ACKNOWLEDGEMENTS. We are grateful to Jim and Theresa Clare for

unpublished information on Chibchanomys and to the late Professor Gustavo

Orcés for support and encouragement during fieldwork. We are indebted to

Robert Voss (American Museum of Natural History) who initially recognised

the uniqueness of the new species and who provided much helpful advice

when commenting on drafts of the manuscript. Particular gratitude is owed to

Darrell Siebert (Natural History Museum) for guidance with the phylogenetic

analysis, and for constructive comments and criticism of the manuscript.

Mark Hafner (Museum of Zoology of Louisiana State University) kindly

provided photographs and measurements of the specimen of C. trichotis from

Peru. As always the staff of the Natural History Museum freely provided

support; in particular we thank Richard Sabin (Mammal Group) and Deryck

Jones (Electronics) for the X-rays, Phil Hurst (Photographic Unit) for the

photographs, while Clive Moncrieff (Biometrics) patiently assisted with the

PAUP analysis.

Fig. 3 Semistrict consensus tree showing hypothetical phylogenetic relationship of Chibchanomys orcesi to other taxa of ichthyomyine rodents.

Consistency Index 0.636, Retention Index 0.786.

128

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— 1906b. A third genus of the Jchthyomys group. Annals and Magazine of natural

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& Carleton, M.D. 1993. A new genus for Hesperomys molitor Winge and

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Meddelelser fra Dansk Naturhistorisk Forening i Kjgbenhavn 5 (3): 20-27.

Bull. nat. Hist. Mus. Lond. (Zool.) 63(2): 129-136

A new species in the asterinid genus Patiriella

(Echinodermata, Asteroidea) from Dhofar,

southern Oman: a temperate taxon in a

tropical locality

POD ~~ > Fr | a

ANDREW C. CAMPBELL _ 0 aaageilh ipscsiie deine

School of Biological Sciences, Queen Mary and Westfield College, University of London, Mile End Road,

London E]1 4NS, UK.

FRANCIS W.E. ROWE

Goldbrook Boarding Kennels and Cattery, Nuttery Vale, Cross Street, Hoxne, Suffolk, IP21 5BB, U.K.

CONTENTS

SSNS) DISS aa ce SAREE Pea o SEBS PEER E CAEL SES CEP EEE eR BRAE SPE MAGEE BRC OPE ER PP SSE NSP eT OPT er SRS See 129

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SYNOPSIS. The status of the hitherto temperate-water asterinid genus Patiriella is briefly discussed, including in its diagnosis

details of actinal plate and ventral/lateral plate arrangement. A new species, P. paradoxa, is described from shallow water on the

Dhofar coast of Oman. The distribution of Patiriella on the Dhofar coast is discussed in relation to the peculiar oceanographic

Issued 28 November 1997

conditions and vicariant events in the region.

INTRODUCTION

Although one of us (ACC) made extensive collections along the

coasts of the Sultanate of Oman between 1983 and 1990 (Campbell

and Morrison, 1988 and Marsh and Campbell, 1991), the echinoderm

fauna of southern Arabia remains incompletely known. However,

those echinoderms which have been recorded are tropical species or

endemic species with tropical affinities (Clark and Rowe, 1971;

Price, 1982; Campbell and Morrison, 1988; Marsh and Campbell,

1991). The discovery (by FWER) of three specimens of a new

species of the predominently temperate-water, asterinid starfish

genus Patiriella Verril (1913) among the Omani shallow water (less

than 10m) collections from two sites in Dhofar therefore poses a

paradox.

Here we briefly discuss the status of the genus Patiriella and

describe fully the new species. This work attempts to explain the

presence of an otherwise temperate-water echinoderm genus in this

tropical location, given the peculiar local oceanic conditions which

include upwelling (see summary by Campbell and Morrison, 1988),

and past vicariant events.

The type specimens have been deposited in the Natural History

Museum, London U.K.

SYSTEMATIC DESCRIPTION

PATIRIELLA Verrill

Patiriella Verrill, 1913: 483; 1914: 263; Fisher, 1919: 410; H.L.

Clark, 1946: 134; Dartnall, 1971: 39; A.M. Clark and Courtman-

Stock, 1976: 80; A.M. Clark, 1983: 365; A.M. Clark and Downey,

1992: 192.

TYPE SPECIES. Asterina (Asteriscus) regularis Verrill, 1870

(1867); by original designation.

DIAGNOSIS

After A.M. Clark in A.M. Clark and Downey (1992), amended.

A genus of Asterinidae with five to about eleven short rays (R up

to c. 60mm); pentagonal to stellate in outline; aborally arched;

primary abactinal plates in two ‘fields’, a slightly irregular radial

(usually the midradial and first dorsal-lateral series on either side)

‘field’ and a regular lateral ‘field’ on either side; mid-radial abactinal

plates with proximal edge trilobed or simply crescentic; the proxi-

mal concave sides of abactinal plates subtend spaces with usually

one to several papular pores, separated by one to several small

secondary plates; abactinal armament comprising relatively few

(<40 per plate) very coarse, almost granuliform, multipillared

130

spinelets; actinal plates distinctly aligned in oblique series between

the second or third, and subsequent adambulacral plates and the

inferomarginal plates and delimiting a membranous, proximal, tri-

angular area which is usually filled by several plates; ventral-lateral

angle of rays supported internally by abactinal plates which meet the

actinal plates by virtue of the oblique alignment of both; towards the

ray base as the ventral-lateral angle becomes less acute with ray

depth, totally internalised plates, spanning between the abactinal

and actinal plates, can be found; actinal armament coarse, short

spines, mostly single, not more than two per plate; furrow spines

usually two (sometimes one) per plate; subambulacral spines one

(rarely two); suboral spines one to four or none; no pedicellariae.

Species included: Patiriella brevispina H.L. Clark, 1938;

Asterias calcar Lamarck, 1816; Asteriscus calcarata Perrier, 1869;

Asteriscus chilensis Lutken, 1859; Asterina dyscrita H.L. Clark,

1923; Asterias exigua Lamarck, 1816; Asterina fimbriata Perrier,

1875; Asterina gunni Gray, 1840; Patiriella inornata Livingstone,

1933; Asterina oliveri Benham, 1911; Patiriella parvivipara

Keough and Dartnall, 1978; Patiriella pseudoexigua Dartnall, 1971

(with subspecies pacifica (Hayashi, 1977, as Asterina); Asterina

(Asteriscus) regularis Verrill, 1870 (1867); Patiriella vivipara

A.C. CAMPBELL AND F.W.E. ROWE

Dartnall, 1969b; Patiriella paradoxa sp. nov.

Other species Patiriella nigra H.L. Clark, 1938 and Patiriella

obscura Dartnall, 1971 are considered to be conspecific with P

oliveri (Benham) and P. pseudoexigua Dartnall respectively by

Rowe (in Rowe and Gates, 1995); Patiriella tangribensis Domantay

and Acosta, 1970, is inadequately described and cannot confidently

be assigned to this genus.

REMARKS

The genus Patiriella Verrill, 1913, has had a rather chequered

history. It has been considered a valid taxon by most recent authors

(e.g. Fisher, 1919; H.L. Clark, 1928; 1938; 1946; Livingstone, 1933;

Madsen, 1956; Dartnall, 1971; A.M. Clark and Rowe, 1971; A.M.

Clark and Courtman-Stock, 1976; A.M. Clark, 1983; A.M. Clark

and Downey, 1992) or a synonym of Asterina Nardo, 1834 (e.g. H.L.

Clark, 1916; 1923; Hayashi, 1940; 1977; Mortensen, 1933 (as a

subgenus of Asterina)). The history and current status of Patiriella

has been most recently discussed by A.M. Clark (1983; 1992 (in

Clark and Downey)) who commented (1992:178) that “The very

coarse and abbreviated, almost granuliform, armament of the upper

side may warrant a supra-specific distinction from A. gibbosa of P.

Fig. 1 a, Scanning electron micrograph of the actinial surface of Asterina gibbosa from Plymouth U.K. Scale bar = 5mm . G = gonopore; b, Scanning

electron micrograph of the actinial surface of ‘Asterina’ cepheus from Wadi Haart, Sadh, Dhofar, Southern Oman. Scale bar = 5mm

A NEW SPECIES OF PATIRIELLA

regularis together with P gunni (Gray) and P. calcar (Lamarck)

from Australia, also P exigua (Lamarck) which extends from the

Indo-West Pacific into the S.E. Atlantic’. She thought reassessment

of the rank of Patiriella was best left to ‘one of several Australasian

specialists to determine . . .’, retaining the species exigua as a mem-

ber of the genus Patiriella. A new review of the family Asterinidae,

including a reappraisal of the status of Patiriella is being undertaken

by F.W.E.R. elsewhere. However, whilst in the present paper we

acknowledge that a very close relationship exists between the genera

Asterina, Patiriella and Patiria (Gray, 1840) (Patiria is considered a

synonym of Asterina by Hayashi (1940) and A.M. Clark (1983 (in

Clark and Downey, 1992)); we accept their separate generic status,

until the matter is more clearly resolved, such a resolution being

outside the scope of this paper '. In taking this stance, we have

amended A.M. Clark’s in Clark and Downey, 1992) diagnosis of

Patiriella to include a description of the alignment of actinal plates

(included by Verrill (1913) in his diagnosis of the genus) and the

internal alignment of actinal/abactinal plates at the ventral/lateral

angle (described by Verrill (1913) as a feature of the family

Asterinidae). As a taxonomic character, actinal plate alignment has

been largely ignored. The exception was Fisher (1917; 1919) who

thought this a useful character when distinguishing his new genus

Paranepanthia (type-species Nepanthia platydisca Fisher, 1913) in

which the plates are aligned obliquely across the actinal surface,

from Asterina in which he saw the actinal plates forming chevrons

across each interradial area, the plates being aligned only parallel to

the furrows along each ray. Fisher (1919) was, however, comparing

Paranepanthia platydisca with the species ‘Asterina’ cepheus (Miiller

& Troschel), and ‘A’. coronata (von Martens) and not the type

species A.gibbosa. In accepting Asterina in this sense, Fisher was

clearly not familiar with the fact that in the type-species, A. gibbosa,

the actinal plates, as in Patiriella and Paranepanthia, are clearly

aligned obliquely between furrow and margin Fig la & b (there is no

question, however, that Paranepanthia is a valid genus (F.W.E.R.).

A.M. Clark (1971, in A.M. Clark and Rowe) noted Fisher’s work

when identifying two specimens of an undetermined species of what

she considered a Paranepanthia from Zanzibar. She considered

(1971:71) the importance of actinal plate arrangement, as a func-

tional and taxonomic character, required investigation. However,

A.M. Clark (1983) did not expand further on this matter in her more

recent revision of the family Asterinidae. Although a recent survey

of the family by one of us (RW.E.R.) does show actinal plate

arrangement to be a useful character in distinguishing some of the

genera, actinal plate arrangement undoubtedly has a functional role

which is probably micro-habitat related. We include description of

the internal structure of the ventral/lateral angle in our diagnosis

following A.M. Clark (1983) who concluded that differences in

arrangement of these internal plates were important in indicating

taxonomic affinities, a conclusion supported herein, though on the

basis also, that it may relate to ray shape within the family.

Patiriella paradoxa sp. nov.

DIAGNOSIS. Stellate species of Patiriella with five rays; abactinal

plates of aboral, radial ‘field’ tri-quadrilobed, forming a delicate

reticulum; papular areas each with three to six papulae and one to

two secondary plates; no suboral spines; proximal actinal plates

each with a single spine.

' Species attributed to Asterina s.s. from outside the Atlantic region are NOT conge-

neric with the type-species, A. gibbosa (Pennant), and require reallocation to other

existing or new asterinid genera according to Rowe (in Rowe and Gates, 1995); the

suggested recognition of Asterinides Verrill (1913) (type-species A. folium (Liitken)) as

a valid subgenus of Asterina, by A.M. Clark (1983; 1992 (in Clark and Downey)) is

supported, but at generic level, by Rowe (in Rowe and Gates, 1995).

131

HOLOTYPE. BMNH (dry) BMNH 1997.1016

TYPE LOCALITY. 0.5 km southeast of Wadi Haart and about 4km

north east of Sadh village, Sultanate of Oman (17° 04'N, 55° 06'B),

intertidal, just below level of the conspicuous barnacle Tetraclita

squamosa rufotincta, collected together with ‘Asterina’ cepheus

(Miiller & Troschel, 1842), A.C. Campbell 4 May 1987 (210050201)

(fig.2).

MATERIAL. In addition to the holotype, two paratypes (dry).

Paratype 1 BMNH 1997.1017, collected at the same locality with the

holotype. Paratype 2 BMNH 1997.1018, Raaha, 2.5km west of Wadi

Ayn, Sultanate of Oman (16° 58'N, 54° 50'E), c. 8m depth, on rocks

amongst corals, coll. A.C. Campbell, 5 Dec., 1986 (210050202)

(fig.2).

EtYMOLOGY. The species name (Lat. paradoxum) refers to the

unexpected occurrence of this predominently temperate-water ge-

nus along an otherwise tropical coastline.

DESCRIPTION OF HOLOTYPE. (Figs 3 a & b; 4a & b). Specimen

stellate in outline, R=17.8mm, r=9.0mm, R/r=1.98; br=10.3mm

(across base of ray between first superomarginal on each side), R/

br=1.73. Orally flat, aborally arched. Rays more or less elongate

triangular, tapering from a relatively wide base to a rounded tip.

Centre of disc delimited by a complete ring of prominent, spinelet-

bearing plates, outside of which a second, more or less complete ring

of less prominent plates evident. The prominent, ovate madreporite

occurs in interradius CD.

Beyond the central disc, the imbricating primary abactinal plates

form two ‘fields’ along the rays. Radial ‘field’ comprising an

irregular, zig-zag series of mid-radial (carinal) plates and the first

dorsal-lateral row on each side. Shape of this ‘field’ elongate-leaf-

shaped along the ray, tapering proximally and distally and widest

at about ’R. Denuded plates deeply notched, mostly quadrilobed

(X-shaped), but, at least between ¥2—4R, some mid-radial plates

tri-lobed (Y-shaped), the plates together forming a relatively deli-

cate reticulum. Papular areas, between the plates, are relatively

large, up to 0.8mm diameter, each subtending 3-6 papulae

between which | or 2 minute, spinelet-bearing secondary plates

usually occur. Primary plates with crystal bodies, except on their

proximal, crescentic ridge which carries 5—7 granuliform spinelets

in a single series; spinelets range from c. 0.30mm long x 0.12mm

wide (straight sided) to about 0.37mm long x 0.14mm wide (these

larger spinelets becoming club-shaped (0.18mm wide) towards

their tip). Lateral ‘field’ comprising about 10 regularly arranged

rows of plates at the base of the rays, closely imbricate with a deep

proximal notch subtending 1-3 papulae occurring in the proximal

half of the first 5 rows of plates, the papulae not extending to the

superomarginal line. Lateral ‘field’ plates with crystal bodies and

from 2-7 spinelets.

Ventral-lateral margin sharply delimited, however neither

inferomarginals nor superomarginals significantly larger than im-

mediately adjacent actinal or abactinal plates respectively. Infero-

marginal plates aligned in the same plane as the actinal surface,

slightly protrude laterally and bear a group of 2—3 minute spinelets.

Superomarginal plates aligned vertically, correspond with the

inferomarginals below them and bear 1—2 spinelets.

About 10 rows of actinal plates counted; the first two extending to

the tip of the ray; plates are aligned both parallel to the furrow but

also distinctly obliquely across the actinal surface between the

adambulacral plates and inferomarginal plates. Oblique alignment

accentuated both by the actinal spination and narrow, shallow

A.C. CAMPBELL AND F.W.E. ROWE

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A NEW SPECIES OF PATIRIELLA

eet

Fig.3 a, Photograph of Patiriella paradoxa, Holotype, from Wadi Haart, Sadh, Dhofar, Southern Oman. Abactinial view. Scale in mm; b, Photograph of

Patiriella paradoxa, Holotype, from Wadi Haart, Sadh, Dhofar, Southern Oman. Actinial view. Scale in mm.

furrows occurring between the oblique lines of plates. First com-

plete oblique line of plates arises opposite the second adambulacral

plate in each furrow, creating a triangular area bounded by the first

adambulacral plates, distal edge of the oral plates and first oblique

series in each actinal intermediate area. This proximal area occupied

by 2 (interradii BC, DE and EA), 3 (interradius AB) or 4 (interradius

CD) plates. A small circular patch of non-calcified skin (up to

0.6mm diameter) occurring adjacent to the distal edge of the oral

plates in interradii AB, CD and EA. Actinal plates each bear a single,

tapering spinelet (up to 0.64mm long x 0.30mm at base) on the

proximal 80% of the actinal surface, the remaining distal plates with

2 small spinelets.

There are 28-29 pairs of adambulacral plates to each furrow.

These are twice as wide as long. First 12—14 plates bear 2 tapering

furrow spines, proximalmost more slender and shorter than distalmost

spine (on adambulacral 3: distalmost spine measures 1.0mm long x

0.35mm wide, at base tapering 0.20mm wide near tip; proximalmost

spine measures 0.78mm long x 0.23mm wide at base, tapering to

0.14mm wide near tip). Proximalmost spine becomes rapidly smaller

and peg-like towards the 12th—14th plate, beyond which the

adambulacral plates each bear a single furrow spine. A single,

stouter, subambulacral spine (on adambulacral 3: 0.87mm long x

0.29mm wide, more or less cylindrical) occurs on each adambulacral

plate. The oral plates each with 4 oral (furrow) spines of which the

apicalmost is longest. No suboral spines.

Gonopores not occurring on the oral surface and cannot be

distinguished aborally.

CoLour. Dried holotype is a uniform, pale ‘museum’ buff colour.

Colour in life is not recorded.

PARATYPE 1. BMNH 1997.1017 has R=19.0mm, r=7.9mm.

R/r=2.4; br=9.4mm, R/br=c.2. Generally very similar in appearance

to the holotype, differing only in minor detail. Two furrow spines

occurring only on the 1st-4th adambulacral plates, thereafter the

plates with a single furrow spine. Small patch of non-calcified skin

occurs adjacent to the distal edge of the oral plates in each interradius,

this proximal actinal triangle being filled by 1 (interradii CD, DE) or

2 (interradii EA, AB, BC) plates.

PARATYPE 2. BMNH 1997.1018 has R=19.0mm, r=10.00mm,

R/r=1.9; br=11.2 mm, R/br=1.7. The dry specimen is contorted. It

differs from the holotype and paratype | in the following features.

Rays are slightly broader at their bases relative to their lengths.

Aborally, centre of disc not delimited by a prominent ring or rings of

plates. Radial ‘field’ of abactinal plates is compact, plates closely

imbricating with only the ridge evident and bearing 5-7 spinelets.

Papular areas contain 1-4 papulae and occasionally a single second-

ary plate. Actinal spinulation coarser, otherwise the plate arrangement

closely similar to the other type specimens. Actinal proximal trian-

gle filled by 1 or 2 plates, no patches of non-calcified skin evident.

Some 36 pairs of adambulacral plates along each furrow; each

bearing a single furrow and single subambulacral spine, with the

exception of the first plate with a second, smaller proximal furrow

spine. Furrow and subambulacral spines flattened along their length

with a squared-off tip. Several furrow spines spaced along length of

the furrow, with bifid tip. Oral plates and spines are similar to those

of both the holotype and paratype 1. The paratype 2 is uniformly

light grey in colour.

HABITAT. The holotype and paratype | were collected on 4.5.87,

0.5km. south east of Wadi Haart and about 4 km. north east of Sadh

village. These were collected, together with ‘Asterina’ cepheus

intertidally on a gently shelving exposed rocky shore with tide pools

just below the level of the conspicous barnacle Tetraclita squamosa

A.C. CAMPBELL AND F.W.E. ROWE

Fig. 4 a, Scanning electron micrograph of the abactinial surface of Patiriella paradoxa from Wadi Haart, Sadh, Dhofar, Southern Oman. Scale bar: 5mm;

b, Scanning electron micrograph of the actinial surface of Patiriella paradoxa from Wadi Haart, Sadh, Dhofar, Southern Oman. Scale bar: 5mm.

rufotincta. During the monsoon period, July-September, the middle

shore is richly cloaked with the green alga Ulva sp. and brown algae

develop on the lower shore. These growths are burnt off by Decem-

ber, and then the intertidal remains almost alga free until the next

south west monsoon. It is worth noting that perennial beds of the

kelp Ecklonia radiata have been recorded off Sadh head (Barratt et

al., 1986) approximately 5 km. to the south west of this site.

Ecklonia may actually occur much closer as fragments were found

washed up on the beach.

Paratype 2 was collected on 5.12.86 at Raaha 2.5 km. west of

Wadi Ayn at 8 m depth on rocks amongst corals. This site faced south

south east and comprised a sandy cove bordered to the east by a ridge

of metamorphic rock sloping down to sand at 10 m. The rock ridge

was well covered with many scleractinian colonies, especially

Acropora sp. During the south west monsoon period this ‘coral

garden’ became completely overgrown with the brown alga

Sargassopsis zanardini. At the time of the collection the S. zanardini

growths had broken up and dispersed. No Ecklonia radiata was seen

growing in the immediate vicinity.

DISTRIBUTION. Known only from the type localities on the coast

of Dhofar, southern Oman.

REMARKS. Based on arrangement and shape of skeletal plates and

their armament, paradoxa is without doubt congeneric with Patiriella

regularis, the type species of Patiriella. It differs from that species,

as with the majority of its congeners in the form of the radial ‘field’

of abactinal plates, the high number of papulae per papular area and

absence of suboral spines. The absence of suboral spines is shared

with four species from the southern Australian coast. Of these P.

parvivipara is a small, precociously viviparous, pentagonal, cush-

ion-shaped star; P. brevispina and gunni have six, non-projecting

rays, and are more or less cushion-shaped and hexagonal in outline.

Additionally, these species differ from P. paradoxa in the increased

frequency of two spines on each of the proximal actinal intermediate

plates, and gunni possesses two subambulacral spines per

adambulacral plate. P paradoxa may appear to be most closely

related to P. calcar, differing most obviously in having a more

delicate abactinal plating and armament and having 5 instead of 7—

11 short rays. Clearly the geographical isolation of paradoxa from

its congeners has resulted in the evolution of a combination of

characters which isolates it within the genus.

A NEW SPECIES OF PATIRIELLA

ZOOGEOGRAPHICAL CONSIDERATIONS.

The genus Patiriella, as accepted herein, is distributed predominently

in temperate seas. No fewer than 10 of the 15 species included occur

in Australian waters (Rowe & Gates, 1995). Six of the species

(brevispina, calcar, gunni, inornata, parvivipara and vivipara) are

endemic to Southern Australia. Two species range across the Tasman

Sea between southeastern Australia and New Zealand (regularis) or

Lord Howe Island and Kermadec Islands (oliveri). One species

(pseudoexigua) is essentially tropical, ranging from northeastern

Australia north to Japan. The somewhat ubiquitous P. exigua ranges

from southern Australia, westward across the southern Indian Ocean

to St. Helena Island off southwestern South Africa in the southern

Atlantic. Of the other species, dyscrita is endemic to South Africa;

calcarata is endemic to Juan Fernandez Islands; chilensis occurs

between Chile and Peru and, fimbriata is distributed from southern

Chile to the Falkland-Magellan area and southern Argentina (A.M.

Clark, 1993). With the exception of fimbriata which has a known

depth range from intertidal to c. 300m, species of Patiriella are

essentially intertidal, occurring at most to about 30m depth (A.M.

Clark, 1993).

The occurrence of a species of Patiriella isolated on the Dhofar

coast of Oman, in the tropical, northwestern Indian Ocean, is

difficult to explain; whether as the result of distributive, accidental or

vicariant events.

To seek explanation by a distributive means requires knowledge

of reproductive strategies within the genus. Unfortunately, we have

been unable to determine reproductive strategy in PR paradoxa

through examination of the gonads, since the three specimens

collected to date were preserved and dried. Since gonopores do not

occur on the oral surface (see p. 133), and it is clearly not viviparous,

then we assume aboral gonopores occur in Pparadoxa and the

strategy involves either planktotrophic or lecithotrophic larvae.

However, life histories of at least seven species of Patiriella occur-

ring in Australia, including P. regularis which also occurs in New

Zealand have been determined (Byrne, 1991; 1992; Byrne and

Barker, 1991). Of these, regularis exhibits an indirect/planktotrophic

developmental pattern with feeding bipinnaria and brachiolaria

larvae; gunni, calcar, pseudoexigua exhibit a direct/lecithotrophic

developmental pattern with planktonic non-feeding brachiolaria

larvae; exigua exhibits direct/lecithotrophic developmental pattern

with a benthic non-feeding brachiolaria larva, vivipara and

parvivipara exhibit a direct/viviparous pattern of development as

intra-ovarian brooders, without larvae. The occurrence on the the

Dhofar coast of Oman of Patiriella originating via larval distribution

from either southern Africa or the Australasian region, even though

the nearest congeners are P. dyscrita (S. Africa) and P. pseudoexigua

(recorded as P. exigua by Koehler, 1910), from the northeastern

Indian Ocean, is difficult to envisage. The present-day water cur-

rents of the Indian Ocean must have been established for at least the

last 15—10my, following separation of the Indian Ocean (see Adams,

1981; van Andel, 1981). The isolation of P paradoxa, on the Dhofar

coast suggests a genetic isolation which is not receiving input from

other parts of the generic range.

Anexplanation of the origin of the temperate-water genus Patiriella

on the tropical Omani coast in association with a secondary agent is,

however, not at first sight, unreasonable to propose. For instance,

Dartnall (1969a) considered the New Zealand species P. regularis

had been introduced into Tasmanian waters amongst oyster spat. The

distribution of P. exigua from southern African to St. Helena Island,

in the south Atlantic, has been attributed, by Mortensen (1933) to

probable transport on the holdfasts of the kelp Ecklonia, which

135

though not established itself on St. Helena was washed ashore there.

A.M. Clark (1992, in A.M. Clark and Downey) considered this a

reasonable supposition since exigua is not recorded from Ascension

Island, to the north of St. Helena where the current is from the east.

The occurrence of Ecklonia radiata, otherwise known only from

around the coasts of South Africa, Australia and New Zealand,

which has been recruited to the southeastern coast of Arabia via the

deep Antartic current from the south (Sheppard, 1992, in Sheppard,

Price and Roberts) might have provided a ‘raft’ for Patiriella to reach

the Dhofar coast, originating either from South Africa or the

Patiriella-species-rich southern coast of Australia. However, such

‘rafting’ or ‘accidental’ introductions tend only to extend species

range, as it has done with Ecklonia. To seek a solution here for

Patiriella requires invocation of the subsequent evolution of the

species P. paradoxa and extinction of its ancestor on the coast of

Dhofar. Once again, the genetic and geographic isolation of P.

paradoxa would not appear to support such an arguement of its

origin on the Dhofar coast, even from its geographically nearest

congeners (see above).

The lack of fossil history of the family Asterinidae does not

preclude ancient history. It may be more productive, therefore to

seek a linkage between the distribution of Patiriella (particularly

considering its very close relationship with Asterina s.s., from the

Atlantic, and Patiria, from the North Pacific) and vicarient events in

the Indian Ocean. The oceanographic and geological configurations

of the area appears to have been relatively stable for at least the last

15-10 my (Powell et al., 1981; Adams 1981).

Coincident with this time the complete opening of Drake’s Pas-

sage (25-15 mya) established the circum-Antarctic current and a

sharp drop occurred in surface and bottom sea-water temperatures

(van Andel, 1981). It can be concurred that the present-day areas of

upwelling, including those within the Indian Ocean were also

established at that time. This is of interest for two reasons. Firstly, a

major area of upwelling is known off the coast of Dhofar (fig. 2).

This produces turbulent, nutrient-rich surface waters and almost

temperate conditions (minimum recorded water temperature of

15.9°C near Sadh; Savidge, et al., 1986) which prevail in the coastal

region of Dhofar for at least 4 months of the year (June—mid

September) (see Currie et al., 1973; Campbell and Morrison, 1988;

Miller and Morris, 1988 for details) and this is coincident with the

known distribution of P paradoxa. Secondly, the general temperate-

water distribution of species of Patiriella (as included herein),

Asterina s.s. (including only species gibbosa, phylactica, stellifera

and possibly pancera; according to FWER, unpublished) and Patiria

(including species miniata, pectinifera and possibly minor; accord-

ing to FWER, unpublished), which appear (with the exceptions of

Patiriella pseudoexigua and Asterina stellifera along the tropical

west coast of Africa, part of its range) not to extend into locations

much, if at all, above the 20°C isotherm (distribution data taken from

A.M. Clark, 1993). The implication of the present-day distribution

pattern of these three genera is that their ancestor was more wide-

spread in cooler parts of the Tethyan system (possibly during the

later part of the Oligocene epoch (c. 40-25 mya) (see van Andel,

1981)) and before its closure, following which event the structural

differences recognised in separating the three genera would appear

to have evolved in the fairly discrete geographical isolation which

occurred between them. Although the distribution of its congener

pseudoexigua across the tropics from northeastern Australia to

southern Japan is somewhat confounding, the isolated, endemic

occurrence of P. paradoxa on the southern Arabian coast might be

explained in terms of its being a relic. This is all the more likely due

to the influence of local seasonal upwelling, providing more temper-

ate conditions, suitable for the genus, at least for part of the year, the

136

species clearly having evolved to survive year-round conditions on

the coast in this tropical part of the world. Upwelling along the west

coast of Africa may well account for the extension of Asterina

stellifera into more tropical regions in the western Atlantic part of its

distribution. It will be of interest now to investigate whether other

shallow temperate-water echinoderms (other than the globally ubiq-

uitous ophiuroid, Amphipholis squamata) (see Price, 1982) or other

invertebrate taxa (see Barratt et al, 1984) might be found to occur in

the Dhofar region, which might support this thesis, particularly since

no other temperate-water distributed taxa appear to have been

recorded from the area.

ACKNOWLEDGEMENTS. A.C.C gratefully acknowledges the diving, field

and laboratory support given by Brian Carnell, Paul Fletcher, Mike Morrison,

Robert Whitcombe, and the Biology Department of Sultan Qaboos Univer-

sity. The field work was financed with a grant from Sultan Qaboos University,

Muscat, Sultanate of Oman.

We also wish to thank two unnamed referees for their valuable and

constructive criticisms of the manuscript. Any remaining errors are the sole

responsibility of the authors.

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Bull. nat. Hist. Mus. Lond. (Zool.) 63(2):137—147

Issued 28 November 1997

Morphological observations on Oncaea

mediterranea (Claus, 1863) (Copepoda,

Poecilostomatoida) with a comparison of Red

Sea and eastern Mediterranean populations

R. BOTTGER-SCHNACK,,

Institut fiir Meereskunde am der Universitat Kiel, Diisternbrooker Weg 20, D-24105 Kiel, Germany

R. HUYS

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5BD, UK.

SYNOPSIS. The taxonomic status of the allegedly cosmopolitan Oncaea mediterranea (Claus, 1863) is reviewed. Comparison

of Claus’ fragmentary original description and the subsequent redescription by Giesbrecht in 1892 revealed significant

differences which might lead to taxonomic confusion. Most authors have adopted Giesbrecht’s identification, but it is clear that

several, as yet unnamed, species have been recorded in the literature under the wrong name O. mediterranea. The species is

redescribed in detail on the basis of material from the Red Sea and eastern Mediterranean. The occurrence of two size morphs in

the eastern Mediterranean populations is briefly discussed. O. mediterranea sensu Bourne (1889) is regarded as species

inquirenda in the genus.

INTRODUCTION

Oncaea mediterranea (Claus, 1863) is one of the most widely

distributed oncaeids (Copepoda: Poecilostomatoida) and according

to the literature assumes a cosmopolitan distribution. It has been

recorded in epi- and mesopelagic layers of all three main oceans

between approximately 60° N and 60° S (Malt, 1983a: Fig. 14;

Heron and Bradford-Grieve, 1995). In some areas such as the

offshore waters of the Lebanon (Malt ef al., 1989) it represents the

most common oncaeid.

O. mediterranea is a well studied species. The naupliar develop-

ment has been documented by Hanaoka (1952), and Malt (1983c)

discussed the polymorphism and pore signature pattern of the

species. Paffenhofer (1993) obtained quantitative information on

reproduction rates and longevity of adult females. It was found that

early copepodids of O. mediterranea ingested about 100% of their

body weight daily when feeding at relatively high food levels.

Average reproduction rates of field-collected females ranged from

5.3 to 13.3 nauplii day~'. According to Paffenhofer (1993) neither

nauplii, nor copepodids or adults of O. mediterranea create a

feeding current, and because of their limited swimming perform-

ance the encounter with food has to be created either by motile food

particles or by the copepod jumping repeatedly to locate a food-rich

environment. O. mediterranea has been observed on discarded

appendicularian houses (e.g. Alldredge, 1972) which probably serve

as a major constituent of detritus and a food source for copepods in

the epi- and mesopelagic zones (Ohtsuka et al., 1996).

Like in many other so-called cosmopolitan planktonic species the

taxonomy of O. mediterranea is potentially confusing. Most identi-

fications of this species are based on the redescription of Giesbrecht

(1892) from the Gulf of Naples, which was excellent by contempo-

rary standards, and do not consider the type description of Claus

(1863, as Antaria mediterranea) from Messina. Comparison of

Claus’ and Giesbrecht’s text and illustrations casts certain doubts, as

to whether both descriptions are based on the same species. As part

of an ongoing study on Red Sea oncaeids the taxonomic history of O.

mediterranea is reviewed and the species is redescribed herein on

the basis of material from the eastern Mediterranean and various

regions in the Red Sea.

METHODS

Oncaeids were collected using a multiple opening-closing net with

a mesh size of 0.05 mm during cruise 5/5 of R/V Meteor in the

Southern and Central Red Sea and in the Eastern Mediterranean Sea.

A station list and sampling data are given in Table 1. The plankton

was initially fixed in 4% formaldehyde-seawater solution buffered

with hexamethylene tetramine and transferred after ca 2 years into a

preservation fluid of 5% propylene glycol, 0.5% propylene

phenoxetol, and 94.5% filtered seawater (Steedman, 1976). Speci-

mens were dissected in lactic acid, and mounted on slides in

lactophenol. All figures have been prepared using a camera lucida

on a Leitz Dialux differential interference contrast microscope.

Total body length and the ratio of prosome to urosome (excluding

caudal rami) were calculated as the sum of the middorsal lengths

Table 1 Station data of sampling with 0.05 mm mesh multiple opening

closing net during R/V Meteor Cruise 5.

Stn. Haul Date Time Latitude Longitude Depthof Total

No. No. (1987) (°N) (°E) Haul(m) water

depth (m)

Red Sea

663 17/4 20.07 Day 22°58.4' 37°19.4' 50-100 1200

703 39/5 03.08 Day 15°34.8' 41°54.9' 0-50 970

39/4 50-100

Eastern Mediterranean

44 31/5 24.01 Day 34°07.1' 31°54.7' 0-50 2530

138

of individual somites measured in lateral view. In the case of

telescoping somites these lengths are measured from the anterior to

the posterior margin. This approach differs from that traditionally

used in oncaeid taxonomy, where the telescoping of somites is not

considered in length measurements. Thus, sizes of the species in this

paper are not directly comparable to those of previous descriptions

(e.g. Heron, 1977; Heron and Bradford-Grieve, 1995) or earlier

studies in the Red Sea (Bottger-Schnack et al., 1989). Length data

given by the latter authors are only about 70% of the length pre-

sented in this paper, due to the excessive telescoping of somites in

the sorting medium.

Descriptive terminology for body and appendages follows that of

Huys and Boxshall (1991). Abbreviations used in the text are: ae =

aesthetasc; CR = caudal rami.

SYSTEMATICS

Order POECILOSTOMATOIDA Thorell, 1859

Family ONCAEIDAE Giesbrecht, 1892

Genus ONCAEA Philippi, 1843

Oncaea mediterranea (Claus, 1863)

(Figs 1-5)

SYNONYMS. Antaria mediterranea Claus, 1863; Oncaea medi-

terranea (Claus, 1863); Oncda mediterranea (Claus, 1863).

ORIGINAL DESCRIPTION. Claus (1863): 159-160, Tafel XXX, Fig.

IRGC), (ED)

OTHER DESCRIPTIONS. Giesbrecht (1892) [as Oncda mediterr-

anea]; Heron (1977) and Heron & Bradford-Grieve (1995) [as

Oncaea mediterranea].

TYPE LOCALITY. ‘Tyrrhenian Sea; Messina.

MATERIAL EXAMINED. (see Table I for locality data)

(1) The Natural History Museum, London: Southern Red Sea: Stn

703; R/V Meteor leg 5/5; collected with MSN 0.05 mm net

(Haul 39/4); depth 50-100 m: 102 Qand 10¢'C'in alcohol

(reg. nos 1996.1095—1114); leg. R. Bottger-Schnack;

(2) Dr Bottger-Schnack (personal collection):

(a) Southern Red Sea: Stn 703; R/V Meteor leg 5/5; collected

with MSN 0.05 mm net (Haul 39/4); depth 50-100 m; leg.

R. Béttger-Schnack: several 2 2 and oO’ Cin alcohol;

(b) Central-Northern Red Sea: Stn 663; R/V Meteor leg 5/5;

collected with MSN 0.05 mm net (Haul 17/4); depth 50—

100 m; leg. R. Bottger-Schnack: several 2 9 and O’C'in

alcohol;

lected with MSN 0.05 mm net (Haul 31/5); depth 50-100

m; leg R. Béttger-Schnack: (i) small form: 1Qand 1

O'dissected on slides, 1 Qin alcohol; (ii) broad form: 2

2 QPdissected on slides, 1 Cin alcohol.

ADULT FEMALE

Body length (measured in lateral aspect; from anterior margin of

rostral area to posterior margin of caudal rami, calculated as sum of

individual somites): 1400 um

Exoskeleton well chitinized. Prosome 2.2 times length of urosome,

excluding caudal rami, 1.9 times urosome length including caudal

R. BOTTGER-SCHNACK AND R. HUYS

rami. Leg 2-bearing somite without conspicuous dorso-posterior

projection (Fig. 1B). Leg 3-bearing somite with conspicuous raised

pore protruding laterally (Fig. 1a). Other integumental pores on

prosome as indicated in Fig. 1A—B. Pleural areas of leg 4-bearing

somite with constricted posterolateral corners.

Proportional lengths (%) of urosomites are 9.7 : 60.7 : 8.2: 8.8:

12.6. Proportional lengths (%) of urosomites and caudal rami are 8.1

SOWA E Ochs Mes) = Os) 2 i>).

Genital double-somite nearly twice as long as maximum width

(measured in dorsal aspect) and twice as long as postgenital somites

combined; largest width measured at anterior third, posterior part

tapering gradually. Paired genital apertures located at about halfway

the distance from anterior margin of genital double-somite; arma-

ture represented by | spine and 1 diminutive spinule (Fig. 1H). Pore

pattern on dorsal surface as indicated in Fig. 1C.

Anal somite 1.3 times wider than long; about 2/3 the length of

caudal rami (Fig. 1C). Secretory pore discernible on either side of

anal opening. Anterior margin of anal opening (vestigial anal oper-

culum) with transverse row of minute denticles. Posterior margin of

somite finely serrate ventrally and laterally (Fig. 1D).

Caudal ramus (Fig. 1F) about 3.0 times as long as wide. Armature

consisting of 6 elements: antero- and posterolateral setae long,

spiniform and unipinnate along medial margin; outer terminal seta

long and plumose; inner terminal seta longest and plumose; terminal

accessory seta more than 2/3 the length of outer terminal seta and

more than twice the length of caudal ramus; dorsal seta about half

the length of terminal accessory seta, plumose and bi-articulate at

base. Inner margin of somite with fringe of long, setules. Dorsal

anterior surface (Fig. 1F) with secretory pore near insertion of seta

II. Dorsal surface covered with numerous small scales (Fig. 1F).

Antennule 6-segmented (Fig. 1E), relative lengths (%) of seg-

ments measured along posterior non-setiferous margin 6.2 : 20.6 :

46.4 : 10.3 : 6.2 : 10.3. Armature formula: 1-[3], 2-[8], 3-[5], 4-

[3+ae], 5-[2+ae], 6-[6+(1+ae)].

Antenna 3-segmented, distinctly reflexed (Fig. 2A). Coxobasis

with row of long, fine spinules or setules along outer and inner

margins and with few additional spinules on proximal and distal part

of outer (exopodal) margin; with bipinnate seta at inner distal corner.

Endopod segments unequal in length; proximal endopod segment

subtriangular forming outer lobate outgrowth bearing spinular patch,

with row of denticles along posterior inner margin. Distal endopod

segment slightly shorter than proximal exopod segment, with nar-

row cylindrical base articulating; with two patches of short spinules

along outer margin; lateral armature consisting of 1 unipinnate

spiniform seta and 3 curved setae, one of them sparsely pinnate, all

armature elements similar in length; distal armature consisting of 5

long unipinnate setae and 2 short naked setae; none of armature

elements spiniform or geniculate.

Labrum (Fig. 2B—C) distinctly bilobed. Distal margin of each

lobe with 4 strong dentiform processes medially, short row of long

fine setules latero-distally and row of minute spinules laterally.

Lobes separated by semicircular vertex covered anteriorly by over-

lapping rows of hyaline petaloid flaps. Posterior part of medial

incision with four rounded integumental thickenings. Anterior sur-

face (Fig. 2B) with short row of spinules either side of median

swelling, raised row of small strong denticles latero-posteriorly;

median swelling with large secretory pore posteriorly. Posterior

surface (Fig. 2C) with group of 3 secretory pores located on proxi-

mal part of each lobe and an additional one basally.

Mandible (Fig. 2D) with minute spinules on surface of coxa;

gnathobase with 5 elements: 1 at subdistal ventral corner, 2 along

distal margin and 2 along subdistal dorsal margin; ventral element

shorter than ventral blade, with long, fine setules along dorsal

MORPHOLOGY OF ONCAEA MEDITERRANEA 139

SSS

“tly

Fig. 1 Oncaea mediterranea (Claus, 1863), 9 (Red Sea). A, habitus, dorsal [a: lateral raised pore, enlarged]; B, same, lateral (appendages omitted), arrow

indicating position of lateral raised pore; C, urosome, dorsal; D, urosome, lateral; E, antennule; F, caudal ramus, dorsal; G, leg 5, dorsal; H, leg 6.

140

R. BOTTGER-SCHNACK AND R. HUYS

Key pt hhh bbb

Fig. 2 Oncaea mediterranea (Claus, 1863), 9 (Red Sea). A, antenna; B, labrum, anterior; C, same, posterior; D, mandible, showing individual elements;

E, maxillule; F, maxilla, [f1, distal seta; £2, proximal seta]; G, maxilliped.

141

MORPHOLOGY OF ONCAEA MEDITERRANEA

Vij

yy)

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Za 52 See

LEE

z 4

Ce eA

SSS

SRO

ZZ;

SSSSS

SSS SS

SOR

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OSS SSNS

ROS

leg 2, anterior [b, terminal part of endopod];

Fig.3 Oncaea mediterranea (Claus, 1863), 9 (Red Sea). A, leg 1, anterior [a, terminal part of endopod]; B

C, leg 3, posterior [c, terminal part of endopod]; D, leg 4, anterior [d, terminal part of endopod].

142

margin; ventral blade strong and spiniform, with row of setules on

posterior surface; dorsal blade strong and broad, with four dentiform

processes around distal margin; dorsal elements setiform, the shorter

hyaline, flat and densely setose, the longer multiipinnate.

Maxillule (Fig. 2E) indistinctly bilobed, with numerous spinules

on anterior and posterior surfaces. Inner lobe subcylindrical, with 3

elements: outermost one spiniform, swollen at base, fringed with

coarse spinules, others setiform and bipinnate; innermost one lo-

cated along concave inner margin at some distance from other

elements. Outer lobe with 4 elements; outermost element spiniform,

curved and unipinnate, longer than the following; other elements

with sparse spinules, innermost one shortest.

Maxilla (Fig. 2F) 2-segmented, comprising syncoxa and allobasis.

Syncoxa unarmed, surface ornamented with 2 spinular rows and 1

large secretory pore. Allobasis produced distally into slightly curved

claw bearing 2 rows of very strong spinules along medial margin;

outer margin with strong seta almost extending to tip of allobasal

claw, ornamented with few strong spinules distally and a thin

hyaline lamella bilaterally, tip of seta with tubular extension (Fig.

2f1); inner margin with slender pinnate seta and strong basally

swollen spine with double row of very strong spinules along the

medial margin and single row of shorter spinules along outer margin

(Fig. 2f2).

Maxilliped (Fig. 2G) 4-segmented, comprising syncoxa, basis

and 2-segmented endopod. Syncoxa unarmed, without surface orna-

mentation. Basis robust, inner margin with 2 spiniform bipinnate

elements nearly equal in length; fringe of long pinnules between

proximal seta and articulation with endopod; two rows of long

setules on anterior surface as illustrated in Fig. 2G. Proximal endopod

segment unarmed. Distal endopod segment drawn out into long

curved claw, with pinnules along proximal half of concave margin;

accessory armature consisting of minute, naked seta on outer prox1-

mal margin and unipectinate spine fused basally to inner proximal

corner of claw.

Swimming legs 1-4 biramous (Fig. 3A—D), with 3-segmented

rami. Spine and setal formulae as follows:

Coxa Basis Endopod Exopod

Leg 1 0-0 1-I 0-1; 0-1; 0,L5 0; I-1; Il,1,4

Leg 2 0-0 1-0 0-1; 0-2; LIL3 1-0; I-1; I,1,5

Leg 3 0-0 1-0 0-1; 0-2; LI1,2 0; I-1; IL,1,5

Leg 4 0-0 1-0 0-1; 0-2; LII,1 0; I-1; IL,15

Intercoxal sclerites well developed, without ornamentation. Coxae

and bases of legs 1-3 with surface ornamentation as shown in Fig.

3A-C. Bases with plumose (legs 1, 4), short naked (leg 2) or very

long naked, outer seta (leg 3); with anterior secretory pore near outer

proximal corner; inner portion slightly produced adaxially in legs 2—

4 (Fig. 3B—D). Inner basal seta on leg 1 long, spiniform and naked.

Respective legs without distinct length differences between rami

with endopod slightly longer than exopod. Bases of spines on

exopodal and endopodal segments surrounded by small spinules

which are most obvious around terminal endopod spines of legs 2

and 4 (Fig. 3b,d). Surface ornamentation of all segments with small

scales, similar to those on caudal ramus (Fig. 1F).

Exopods. Outer margin of exopod segments with well developed

serrated hyaline lamella; inner margin of proximal exopod segments

with long setules. Secretory pore present on posterior surface of

distal segments. Hyaline lamellae on outer exopodal spines well

developed. Terminal spine of leg 1 equal in length to (leg 1) or

shorter than (legs 2-4) distal exopod segment.

R. BOTTGER-SCHNACK AND R. HUYS

Endopods. Outer margin of endopod segments with fringe of long

setules. Inner seta of proximal endopod segment slightly swollen.

Distal endopod segments with single secretory pore on posterior

surface; distal margin of legs 2 and 3 produced into conical projec-

tion ornamented with minute spinules anteriorly (Fig. 3B,C,b,c).

Outer subdistal spine equal in length to distal spine in legs 2-4.

Outer margin of distal segment of leg 1 terminating in a long process

obscuring insertion of distalmost inner seta (Fig. 3a). Inner setae of

distal endopod segments of legs 2-4 with spinule comb along

proximal inner margin; this comb less obvious in leg 1; also present

on distal inner seta of middle endopod segment in leg 4.

Leg 5 (Fig. 1G) comprising small plumose seta arising from

lateral surface of somite, and small free segment representing exopod.

Exopod slightly longer than wide, bearing stout curved seta apically

and smaller slender seta inserted subdistally; surface covered with

minute denticles

Leg 6 (Fig. 1C, H) represented by operculum closing off each

genital aperture; armed with a spine and a short spinule.

Egg-sacs paired, oval-shaped; each sac containing approximately

40 eggs (diameter 50-65 pm).

ADULT MALE

Body length (measured in lateral aspect; from anterior margin of

rostral area to posterior margin of caudal rami, calculated as sum of

individual somites): 960 um. Sexual dimorphism in antennule,

antenna, maxilliped, legs 5—6, caudal ramus and in genital segmen-

tation.

Proportional lengths (%) of urosomites (excluding caudal rami)

9.4 : 70.0.: 3.4 : 3.8 : 3.0: 10.5 ; proportional lengths (%) of

urosomites (caudal rami included) 8.4 : 62.1 :3.0:3.3:2.7:8.1:

11.6. Leg 5-bearing somite with transverse row of denticles dorsally

(Fig. 4D). Caudal rami 1.8 times longer than wide, markedly shorther

than in female. Caudal setae with proportional lengths as in female.

Dorsal surface of genital somite covered with scale-like structures in

anterior half, arranged in a specific pattern (not figured). Surface of

genital flaps and ventral surface of anal segment ornamented with

several rows of small spinules (Fig. 4E). Dorsal surface of caudal

ramus covered with minute scales as in female.

Antennule (Fig. 4B) 4-segmented; distal segment corresponding

to fused segments 4—6 of female; relative lengths (%) of segments

measured along posterior non-setiferous margin 14.5 : 19.7: 42.1:

23.7. Armature formula: 1-[3], 2-[8], 3-[4], 4-[11+2ae+(1+ae)].

Antenna as in female, except for lateral armature on distal endopod

segment (Fig. 4G), with third spiniform element being much stouter

than in female and shorter than the 2 adjacent setae.

Maxilliped (Fig. 4C) 3-segmented, comprising syncoxa, basis

and 1-segmented endopod. Syncoxa without surface ornamentation,

unarmed. Basis robust, particularly inflated in proximal half form-

ing bulbous swelling; anterior surface with 2—3 transverse spinular

rows in addition to row of short flat spinules along inner margin

(Fig.4C); posterior surface with 3 rows of short spatulated spinules

of graduated length along palmar margin (Fig. 4C); with 2 small

naked setae inserted within the longitudinal cleft, the proximal one

being slightly longer than the distal one. Endopod drawn out into

long curved claw, concave margin unornamented; accessory arma-

ture consisting of short, unipectinate spine basally fused to inner

proximal corner of claw; claw with minute hyaline apex (Fig. 4c).

Legs 1-4 with armature as in female; variable number of spinules

on conical projection of endopods of legs 2 and 3.

Leg 5 (Fig. 4D, F) exopod not delimited from somite, general

shape and armature as in female; small plumose seta arising from

lateral surface of somite as in female.

Leg 6 (Fig. 4E) represented by posterolateral flap closing off

MORPHOLOGY OF ONCAEA MEDITERRANEA 143

Se—_

C4 ej

_——

SSS

SLOSS.

SSS

Fig.4 Oncaea mediterranea (Claus, 1863), (Red Sea). A, habitus, dorsal; B, antennule; C, maxilliped, anterior [c, tip of claw, showing hyaline lamella];

D, urosome, dorsal; E, urosome, ventral; F, same, lateral (spermatophores immature); G, antenna, distal endopod segment.

144

genital aperture on either side; covered by pattern of denticles as

shown in Fig. 4E; posterolateral corners protruding laterally so that

they are discernible in dorsal aspect (Fig. 4D).

Spermatophore oval, of variable size according to state of matu-

rity (Fig. 4F); swelling of spermatophore during development not

affecting shape or relative size of genital somite.

TAXONOMIC HISTORY

Claus (1863:159) presented a concise original description of Antaria

mediterranea with some illustrations of both sexes. Later Claus

(1866) identified two size groups upon re-examination of original

material from Messina: the large one [1.3 mm excluding CR setae]

being the one that was mentioned in the original description; the

small one [0.8—0.9 mm] which he also recorded from Nice; no

overlap was found between both size morphs from Messina.

Lilljeborg (1875) recorded a single specimen from Mosselbay

(Spitsbergen), and Car (1884) found it in plankton samples taken off

Trieste in the Adriatic Sea. Both authors, however, gave no descrip-

tions or figures and their records therefore remain unconfirmed.

Bourne (1889) [as Oncaea mediterranea] found 2 females in

surface waters off Plymouth. Giesbrecht (1892) regarded this record

doubtful and pointed out the discrepancy between the text and

figures with regard to the CR/anal somite length ratio. Careful

comparison of this character is hampered by the fact that Claus

(1863) only figured the female in lateral aspect and by the possibility

that the urosome is considerably telescoped in the specimen illustrated

by Bourne. Giesbrecht’s statement that Bourne’s and Claus’ speci-

mens differ fundamentally in maxilliped structure is equivocal since

he compared the former with what he had identified as O.

mediterranea rather than with the original type material. On the

basis of Brady’s (1883) illustrations [Challenger Expedition], Bourne

(1889) also tentatively regarded Onc@a obtusa (Dana) as a possible

synonym of O. mediterranea, however, as Giesbrecht (1892) and

subsequent authors suspected, the latter could as well be conspecific

with O. venusta Philippi.

Giesbrecht (1892) reviewed the earlier literature on Antaria and

Oncaea, and summarized the synonymies of the respective species

known at that time. Dana’s (1849) species Antaria obtusa and A.

crassimana were included under the synonymy of both O. venusta

and O. mediterranea, reflecting the author’s undecisiveness on this

matter. Giesbrecht redescribed O.-mediterranea on the basis of

material from Naples and distinguished two colour varieties flava

and rubra.

Comparison of Giesbrecht’s illustrations with Claus’ original

description, however, raises some doubts as to the conspecificity of

the Messina and Naples specimens. The major obstacle in this

comparison lies in the form, position and size of the setae on the

maxillipedal basis which is usually considered as an important

discriminant in oncaeid systematics. The issue is even more compli-

cated by the discrepancy between text and figures in Claus’ original

description. Claus (1863) stated that there are two ornated setae on

the palmar margin of the basis, yet in his figure (Tafel XXX, Fig. 6)

only one naked seta is illustrated. From the position of this seta,

being located halfway the inner margin, it is conceivable that Claus

has overlooked the proximal seta. This hints at the possibility that

Claus’ O. mediterranea is related to the ‘englishi species-group’

which includes O. ornata Giesbrecht, 1891, O. shmelevi Gordejeva,

1972, O. englishi Heron, 1977 and O. alboranica Shmeleva, 1979.

In these species the distal element is long, slender and minutely

pinnate, whereas the proximal one is spiniform and because of its

R. BOTTGER-SCHNACK AND R. HUYS

small size easily overlooked or misinterpreted as a spinule. In

Giesbrecht’s O. mediterranea, however, both elements are (1) of

about the same size and only half the length of the proximal seta in

the Messina material, and (2) positioned differently, i.e. the proxi-

mal one at 1/3 distance from the syncoxa-basis joint, the distal one

at 2/3 distance. A second possibility is that the long palmar seta in

Claus’ original illustration is in fact a maxillary element superim-

posed on the maxilliped since Claus believed that both appendages

represented the rami of a single limb, i.e. the maxilliped.

Other differences are found in the female leg 5 which is longer in

the original description and the shape of the genital double-somite in

lateral aspect which does not have the pronounced swelling anterio-

ventrally as shown in Giesbrecht’s illustration (Taf. 4, Fig. 16). The

male of Claus’ O. mediterranea shows an exceptionally long leg 5

exopod (his Taf. XXX Fig. 7) which might or might not be free. This

character has thus far been found in only a small number of Oncaea

species belonging to the notopus group, such as O. damkaeri Heron,

1977 and O. parila Heron, 1977 (Heron, 1977; Heron et al., 1984)

which also display a very long leg 5 exopod in the females. Species

of thenotopus group have a setation pattern on the female maxilliped

which is significantly different from that displayed in the englishi

group which raises the suspicion that Claus (1863) might well have

based males and females on different species.

As a result of this comparison it is clear that Claus’ original text

and drawings contain several internal inconsistencies and lack the

detail that is necessary to allow unequivocal identification. The

setation of the maxilliped is a potentially critical character in this

process as confirmation or refutal of Giesbrecht’s identification

depends on whether more weight is given to the text statement or to

the illustration. Given the fact that Claus’ figures of the other

cephalic appendages are similarly poor (setation elements are miss-

ing from every limb) it is preferred here to give more weight to the

text as this will lead to nomenclatural stability. Pending the rediscov-

ery of Claus’ types (which are in all probability lost) this admittedly

subjective decision is the best course of action. In view of the grossly

fragmentary original description in which the sexes were based on

two different — but unidentifiable — species and in the absence of

formal holotype designation the taxonomic problem is in our opin-

ion unsolvable. Moreover, it is considered highly unlikely that

collection of topotype material from Messina would be informative

as 130 years have lapsed since Claus’ discovery of the species in an

open pelagic environment that might have been subjected to major

changes since, such as the opening of the Suez Canal in 1869.

O. mediterranea (Claus, 1863) sensu Bourne (1889) is clearly

different from the Mediterranean material and is regarded here as

species inquirenda in the genus.

Other records of O. mediterranea.

O. mediterranea has been recorded from a wide range of localities

such as the Antarctic (Heron, 1977) and the Red Sea (Bottger-

Schnack, 1988). Many of its records, however, remain unconfirmed

such as the Red Sea records of Cleve (1900, 1903) and Thompson

and Scott (1903) [compiled by Halim (1969)]. Since most authors

have followed Giesbrecht’s identification and ignored Claus’ origi-

nal description it is likely that at least one, as yet unnamed, species

became established in the literature under the wrong name O.

mediterranea. For example, re-examination of material collected

during the Terra Nova and Challenger expeditions (deposited in The

Natural History Museum) proved to belong to at least two distinct

species differening in several aspects from O. mediterranea. Scott

(1894) recorded this species (‘1 or 2 females’) from the Gulf of

Guinea, but re-examination of his illustrations leave little doubt that

MORPHOLOGY OF ONCAEA MEDITERRANEA

UE LLL : E

SSNS

<SS

LEE

SSS

TIN

N Ik

HIN

AB +; 200um

145

50 um

Fig.5 Oncaea mediterranea (Claus, 1863), 2 , broad form (eastern Mediterranean). A, habitus, dorsal; B, same, lateral; C, caudal ramus.

he was dealing with a species of the englishi group. Supporting

evidence for this is found in the morphology of the antenna,

maxilliped and leg 4. Dakin and Colefax (1940) recorded the species

from New South Wales from a depth of 0-200 m were it was ‘rare’

but it is doubtful whether they were dealing with O. mediterranea

since material collected from the Great Barrier Reef and identified

with Claus’ species proved upon re-examination to belong to a

closely related undescribed species. Razouls (1974) described O.

mediterranea from the region of Banyuls-sur-mer (Golfe du Lion)

and summarized previous records from the Mediterranean Sea. His

drawing (Fig. 4G) of the male shows a terminal conical projection on

the endopod of swimming leg 4 which disproves his identification as

this is a character exclusively found in representatives of the conifera-

similis complex. Ferrari (1973, 1975) recorded the species from the

Gulf of Mexico and the northern Caribbean Sea and distinguished

three size variants, the status of which will be discussed below under

‘form variants’ of O. mediterranea. Boxshall (1977a) recorded O.

mediterranea from the Northeastern Atlantic near the Cape Verde

Islands and in another report (Boxshall, 1977b) gave detailed infor-

mation on their vertical distribution and diurnal vertical migration.

146

Many more records of O. mediterranea from different locali-

ties in the world ocean are known [see Malt (1983a) for a

review], but are not considered here, because they did not

include figures or a description that positively identified the

species.

FORM VARIANTS

In the eastern Mediterranean, O. mediterranea exhibits two

forms in both sexes: a smaller and more slender form and a

larger, more robust form. These forms differ only in the general

appearance and in the length : width ratio of the caudal ramus in

the females. The difference in general habitus between the two is

caused by the greater width of the leg 2-bearing and leg 3-

bearing somites in the larger form (Fig. 5A). In the females, the

length to width ratio of the caudal rami is larger in the broad form

(4:1) (Fig. 5C) as compared to the small form (3:1, cf. Fig. 1F),

whereas in the males no corresponding difference could be

found. No other morphological differences were discernible

between the two morphs. The small form of O. mediterranea

corresponds to the small variant described by Malt (1983a),

while the large form seems to be similar to her ‘typical’

mediterranea form. Only one size morph of O. mediterranea

was found in the Red Sea, which can be identified as the small

form found in the eastern Mediterranean and on the basis of the

length to width ratio of caudal rami seems to resemble the small

form of O. mediterranea recognized by Malt (1983b). A small

colourless form of O. mediterranea (in addition to the larger

‘orange-golden’ form) was found south of New Zealand by

Farran (1929) but no further description was provided.

Ferrari (1975) reported the existence of three distinct size

groups in O. mediterranea of the Gulf of Mexico. The two larger

forms differed only in size, but had a similar length to width ratio

of the caudal ramus of 4:1. On the basis of their morphometry

they might be assigned to the typical form of O. mediterranea

(sensu Malt). The smaller size group had a length to width ratio

of the caudal ramus of 3.0—3.5:1 and might thus be similar to the

small form sensu Malt and to that found in the Red Sea. The

smaller morph was initially regarded as a potential new species

(Ferrari, 1973) since it differed both in the proportional lengths

of body somites as well as in total body size. Subsequently,

Ferrari (1974), quoting O. conifera Giesbrecht, 1891 as an

exemplar of gross relative size variation in oncaeids, considered

it as merely another size group of O. mediterranea since not

structural differences could be detected in the appendages. The

recent outstanding work on the conifera-complex by Heron and

Bradford-Grieve (1995) has amply demonstrated that much of

the ‘variation’ in O. conifera can be explained by the fact that

morphologically similar species can co-exist and that this mor-

phological similarity can even cause anomalies in the mate

guarding configurations of certain species. It is conceivable that

many of the form variants represent genuine species which can

no longer be discriminated using traditional o-taxonomical

methods. For example, Malt (1983c) mapped the pore signature

pattern of the two female forms of O. mediterranea from the

North Atlantic but failed to reveal any significant difference

between them. It seems therefore that the question whether the

large morph found in the Eastern Mediterranean represents a

sibling species of O. mediterranea can perhaps only be resolved

by breeding experiments or alternative methods using molecular

data such as enzyme electrophoresis, immunological distance

methods or ribosomal RNA sequencing.

R. BOTTGER-SCHNACK AND R. HUYS

ECOLOGICAL NOTES

Geographical distribution.

O. mediterranea is distributed throughout the Red Sea (see review in

Halim (1969); Bottger-Schnack, 1990b, 1995). It was also found in

small mesh net samples from the northernmost part of the Gulf of

Aqaba (Bottger-Schnack, unpubl.).

Vertical distribution and vertical migration.

During summer and autumn, when a strong seasonal thermocline is

developed, the core of the O. mediterranea population during the

day is situated in the zone below the thermocline, at 50-150 m

(Bottger-Schnack, 1990a, unpubl.). Parts of the population migrate

into the upper 50 m during the night, with males showing a greater

proportion of migrating individuals than females. For females, a

bimodal vertical distribution can be found during these seasons, with

the lower part of the population dwelling in the 250-400 m layer, in

the core of the oxygen minimum zone. In the northern Red Sea the

mesopelagic populations of females are found at greater depths than

in the central area, corresponding to the regional differences in the

depth of minimal oxygen concentrations (Bottger-Schnack, 1990b).

These deep dwelling populations are not found during winter

(Bottger-Schnack, 1990b).

Seasonal variation in abundance (central Red Sea).

No consistent seasonal variation in abundance was found for O.

mediterranea in the central Red Sea, indicating that the populations

are not substantially recruited by those from the south (Bottger-

Schnack, 1995).

ACKNOWLEDGEMENTS. We wish to thank Prof. J. Lenz, Prof. D. Schnack

and Dr H. Weikert for support and help. Dr G.A. Boxshall (NHM) and Dr ED.

Ferrari (Smithsonian Institution) reviewed earlier drafts of the manuscript.

This study was supported by a Deutsche Forschungsgemeinschaft grant Le

232/18-1.

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